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Established in 1962
Edited by WILLIAM HOVANITZ
Volume 1
1968
Published at
1160 W. Orange Grove., Arcadia, California, U.S.A.
THE JOURNAL OF RESEARCH
ON THE LEPIJOOFTlRA\
CONTENTS
Volume 7 Number 1
March, 1968
Controlled environment experiments with
Precis octavia Cram. L. McLeod 1
Ecological and Distributional notes on Erebia disa
in central Canada John H. Masters 9
Methods for Studying the Chromosomes of Lepidoptera
Thomas C. Emmel 23
Note on Vital Staining of Actias luna silk
John M. Kolyer 29
Present and Ice Age Life Zones and Distributions
W. Hovanitz 31
Trials of several density estimators on a butterfly
population W. R. Hanson and W. Hovanitz 35
Habitat — Ar^tjnnis callippe laurirui
W. Hovanitz 50
On Mexican Satyridae Lee D. Miller 51
Habitat — Pieris beckeri W. Hovanitz 56
Identity of the moth “Stretchui” behrermana
with new synonymy. J. S. Buckett 57
Volume 7 Number 2 June, 1968
Studies on Xearctic Eucliloe
Part 5. Distribution Paul Opler 65
Species in the Cenera Polui and Euxoa
John S. Bnek('tl S7
Variation in Ciolor and Maculation
in Nemarid pidcherrima
John S. Buckett and T. A. Sears 95
A New Subspecies of Callophrys dumetorum
C. A. Goreliek 99
Note on Damaged Specimens
John M. Kolyer 105
The Generic, Specific and Lower
Category Names of Nearctic Butterflies.
Part 7. The Genus Dryadula
Paddy McHenry
112
Field Studies of Catocala Behavior
Ronald R. Keiper
113
Habitat: General Type Locality,
Glaucopsi/che hjf^damus xerxes
Plcbcjus icariodcs phercs
W. Hovanitz
122
Life History of Satijrium sylvinas dryopc
T. C. Emmel and J. F. Emmel
123
Habitat: Specific type locality,
Plebcjus icariodes missionensis
126
The Generic, Specific and Lower Category
Names of Nearctic Butterflies.
Part 8. The Genus Agrmdis Paddy McHenry 127
Daytime vision by the moth, Exyra
ridin<isi Vernon M. Kirk 131
Volume 7 Number 3 September, 1968
Population Structure of Oeneis melissa semidea
C. S. Anthony 133
A Hybrid Limenitis from New York
Arthur M. Shapiro and James D. Biggs 149
Population Biology of the Neotropical Satyrid
butterfly, Euptychia hermes:
1. Interpopulation movement, etc.
Thomas C. Emmel 153
Rearing technique for speeding up larval stages
Noel McFarland 166
Habitat: Euphydryas editha wrighti
Fred Thorne 167
Volume 7 Number 4 December, 1968
Population of Danaus plexippus
in Southern Calfornia
F. A. Urquhart, N. R. Urquhart, and F. Munger 169
Habitat: Zerene caesonia eurydice
W. Hovanitz 182
The Effect of Pterin Pigments on Wing
Coloration of Four Species of Pieridae \
Edward J. Pfeiler, Jr. 183
Hilltopping as a Mating Mechanism
to Aid the Survival of Low Density Species
James A. Scott
191
THE JOURNAL
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M I I )(S
published by
The Lepidoptera Research Foundation, Inc.
at
1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz
Associate Editors:
Thomas C, Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601.
Maria Etcheverry, Centro de Estiidios Entomologicos, Casilla 147, Santiago,
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T. N. Freeman, Div. of Entomology, Dept, of Agriculture, Ottawa, Ontario,
Canada.
Brian O. C. Gardner, 18 Chesterton Hall Crescent, Cambridge, England.
G. de Lattin, Zoologisehes Institut, Universitat des Saarlandes, Germany.
Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.
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yt'ar, Spring (March), Summer (June), Autumn (September), and Winter (December)
by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
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Journal of Research on the Lepidoptera
7(1): M8, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
CONTROLLED ENVIRONMENT EXPERIMENTS
WITH PRECIS OCTAVIA CRAM.
(NYMPHALIDAE)
L. McLEOD, B.Sc., F.R.E.S.
25 Sleford Close, Balsham, Cambridgeshire, England
INTRODUCTION
“Seasonal Dimorphism” has long been of great interest to
lepidopterists because the insects exhibiting this phenomenon
did not conform with the early accepted principles of systemat-
ics which were based on pigmentation. Only in the early years
of this century were certain previously distinct species shown to
be two extreme forms of the same “seasonally dimorphic” spe-
cies. This was the case with Precis octavia Cramer, an African
butterfly. As a result of such examples, the principles of system-
atics were revised and based on taxonomic differences of micro-
scopic organs instead of on pigmentation alone.
Since then, many studies have been made of the palearctic
“seasonally dimorphic” species but little experimentation has
been carried out on tropical species.
The tropical genus Precis undoubtedly exhibits the most
striking differences between seasonal forms. The characters
generally affected are size, wing shape (especially in tailed
species) and wing colour and markings of both surfaces.
Two races of Precis octavia occur in Africa. The northwestern
race octavia distributed from Sierra Leone, Congo, Ethiopia, to
Somalia, and the southern race sesamus ranging from Angola,
Kenya, Rhodesia to the Cape of South Africa. In the southern
race, form natalensis Staudinger is predominantly red in colour
on both surfaces and form sesamus Trimen is predominantly blue
on the upperside and dark brown /black on the underside.
1
2
McLEOD
J. Res. Lepid.
TABLE_1
METEOROLOGICAL DATA
KAREN. NAIROBI AREA
MONTH
Rain
inches
( Mean)
Temperature
«C
( Mean )
Max.
®C
(Mean)
Min.
®C
( Mean)
Mean
Hours
Sun
per
day
Mean
Sun
per
day
Relative
Humidity
8.30-14.30
JANUARY
1.47
20.2
25.6
15.0
9.7
40.4
57
FEBRUARY
2.13
21 . 1
27.8
6.1
9.6
40.0
56
MARCH
5.27
21 . 1
28.9
10.0
8.3
34,6
61
APRIL
7.71
20.7
28.3
11.1
7. 1
29.6
69
MAY
5.17
19.6
27.8
10.6
6.3
26.3
71
JUNE
1.62
18.0
27.8
5.6
4.5
18.8
69
JULY
0.59
17.1
25.6
5.0
4.1
17.1
68
AUGUST
0.97
17.4
25.6
4.4
4.4
18,3
67
SEPTEMBER
0.92
19.0
27.8
5.6
6.5
27.1
59
OCTOBER
1.93
20.1
27.8
7.8
7.4
30,8
61
NOVEMBER
4.02
20.0
24.4
12.2
7.2
30.0
66
DECEMBER
2.49
19.7
25.6
11.1
8.5
35.4
63
Total
34.29
19.5
Mean
1964-1965
Kabete
Karen
Met. Stn,
Mean 39
years
Minimum Temperature recorded at Karen 3.9®C
7(1): 1-18, 1968
PRECIS OCTAVIA
3
The southern race F. octavia sesamus is locally common in
Kenya. In the Karen area, fifteen miles from Nairobi, the butter-
flies may be seen flying along the edges of the Ngong Forest
and in the late afternoon can often be seen assembling under
the eves of houses and in small stone quarries where they shelter
for the night.
Larval stages occur throughout the Ngong, Karen and Kikuyu
aeras of Kenya where Coleus forskohlii, the food plant, is used
by African smallholders as a hedge plant.
The majority of butterflies seen are either of form natalemis
or of form sesamus. Intermediates can sometimes be taken but
these are uncommon (Butler 1901, Clarke & Dickson 1953,
Pinhey 1949).
Two generations normally occur in a year. During September
to November f. sesamus is in the majority and from January to
April f. natalemis, but there is a normal overlap of survivors.
NATURAL ENVIRONMENTAL CONDITIONS
Although Karen is only 90 miles south of the equator, it is at
an altitude of 6000 ft. and temperatures are moderate. The dur-
ation of daylight is a fairly constant 12 hours per day.
In Table 1 can be seen meteorological data for the Karen area.
The information is quoted here as being typical of an area of
Kenya in which P. octavia sesamus is found in both of its season-
al forms.
It is not at first obvious from the information on Table 1 that
there is a “cold” season and a “warm” season in this area. There
is little difference between the roaximum temperatures for each
month and also little difference between the minimum tempera-
tures. These figures are somewhat misleading because during
the period June to August, the higher temperatures are only
maintained for a short time each day. During the period Decem-
ber to January the higher temperatures are maintained for the
majority of the day and low temperatures are only achieved
for very short periods. It is for this reason that I have included
figures of mean daily sunlight duriation and also expressed sun-
light as a percentage.
Kenya has two rainy seasons, the “short rains” during October
to November, and the “long rains” of March to May. The mean
monthly daytime relative humidity varies little. When one con-
siders that the larvae feed on the flush of vegetation which oc-
curs at the end of, and immediately after the rains, it can be
seen that the late instars and pupae of the two generations ex-
Parent - Vild V f, natalensis ( July/Aug. 1963)
4
McLEOD
/. Res. Lepid.
Complete mortality resulting from extreme temperature
7(1): 1-18, 1968
PRECIS OCTAVIA
5
perience the two differences in temperature and light intensity
mentioned here.
The above information tends to rule out humidity as a factor
concerned with the production of the two colour forms of this
butterfly. I decided, nevertheless, to include humidity as one of
the variable factors in a number of simple experiments.
EXPERIMENTAL
Towards the beginning of this century, Dorfmeister, Merri-
field, Standfuss, Suffert and Weismann showed that in the family
Nymphalidae, heat causes light colouration and cold causes dark
colouration.
Experiments were performed on P. octavia by Marshall (1902)
but definite conclusions were not forthcoming. Marshall sug-
gested that the two extreme forms always alternate with each-
other and that intermediate forms can be produced by shock
treatment or abnormal conditions. (Rothschild 1918). Clark &
Dickson ( 1957 ) in their study of the life cycle of P. octavia, did
perform experiments but reported that f. sesamus was not pro-
duced under the warmer conditions of Durban.
During 1965, 1966 and 1967, I reared numbers of P. octavia
sesamus under controlled environmental conditions in three dif-
ferent laboratories.
Food Supply I had previously shown with English species of
Pieridae and Nymphalidae that supply of food only affects the
size of the imago. Prior to these experiments, larvae of Precis
archesia Cram, and P. octavia were starved during their fifth
instar and specimens of only. 30 mm. were produced. Larvae
which were given a plentiful supply of fresh food throughout
their larval life produced specimens which averaged 50 mm.
Consequently fresh supplies of food were provided daily in
these experiments.
Light Observations on larvae in the field showed that they
tend to keep to the shadier sections of their food plant and avoid
strong sunlight. This is not surprising because temperatures in
direct sunlight are very high in Karen. Light intensity is not
likely therefore to be a factor concerned in the production of
seasonal forms. However, other research workers have suggested
that duration of light may play an important part in the produc-
tion of seasonal forms of Araschnia levana L. of Europe. Despite
the fact that the duration of daylight is a constant 12 hours per
day throughout the year in East Africa where both extreme
forms occur, I decided to include three variables of light. These
were zero light, 12 hours per day, and constant light.
Precis octavia (Cram)
6
McLEOD
/. Res. Lepid.
r-
v£)
(T-
O
0
1
kD
vC
CT'
Complete mortality resulting from extreme temperature
7 (1): 1-18, 1968
PRECIS OCTAVIA
7
Temperature & Humidity Earlier experiments were perform-
ed in cabinets which lacked modern facilities. Humidity was
maintained at a low level using crystals of silica gel which were
replaced daily. High humidity was maintained using water
bottles with cotton wool wicks. Only the higher temperatures
could be maintained at a constant level using electrical heating.
Later, certain of the experiments were performed in modern
constant environment rooms. Here, the control of humidity and
temperature was facilitated by modern humidifying, cooling and
heating equipment.
In all experiments the egg and first instar larvae were kept
under moderate conditions, and larvae were introduced into
more extreme conditions at the second instar. This procedure
was found to greatly reduce early mortality.
Numbers of individuals reaching maturity were not as high
as I intended. Mortalities from disease were often severe espe-
cially under conditions of high humidity, and on two occasions
all culture insects were completely destroyed by safari ants
Donjlus nigricans Illig. This drastically reduced the numbers
of insects available for the experiments and made a properly
replicated trial impossible.
RESULTS
Extreme variation in pigmentation was recorded in larval,
pupal and adult stages.
Throughout the experiments the majority of larvae exhibited
five instars. In one instance however, several larvae in the same
cage exhibited seven instars. The cage concerned was maintain-
ed at a temperature of 21 °C with no humidity control and in
complete darkness. The reason for the occurrence of the extra
instars is not known. Clarke & Dickson (1953) also record the
occurrence of five, six and seven instar larvae and suggest that
it forms a mechanism for staggering emergence of adults.
Head capsules were collected anl mounted from each cage
and these completely verify the observations.
Pigmentation of Larvae
Early instars did not vary their pigmentation according to the
differing environmental conditions. It soon became apparent
that the colour of the fifth instar larvae varied according to tem-
perature. At the lower temperatures the larvae were black and at
the higher temperatures they were orange, without exception.
The larvae which exhibited seven instars showed variations in
pigmentation in the seventh instar only.
8
McLEOD
J. Res. Lepid.
Figs. 1-6 Precis octavia Cram.
Upperside, left; underside, right. 1 and 2, f. natalensis Staudinger ^ ;
3 and 4, f. transiens Wichgraf $ ; 5 and 6, f. transiens Wichgraf $ .
7(1): 1-18, 1968
PRECIS OCTAVIA
9
The situation was further complicated by the discovery that
there were two strains of larvae which exhibited different colour
forms. The earlier experiments of 1965 (Table 2) produced all
'plain” larvae and the experiments of 1966-67 (Table 3) pro-
duced a mixture of “plain” and “striped” larvae. The appearance
of a second type of larva in the second series of experiments
corresponded with the introduction of another strain of adult
(derived from the same locality).
The two types of larvae and the way in which the pigmenta-
tion of the final instars is changed with temperature, are de-
scribed here.
Plain Larvae — Final Instar
21 °C. and below
Almost entirely velvety black with metallic blue bases to the
spines. Two yellowish patches occur in each thoracic segment,
situated one on each side of the mid-dorsal line. These patches
may be absent at low temperatures.
24 °C.
As above but the yellowish patches extend the entire length
of the larva along the mid-dorsal line and the lateral ridge.
27-32 °C.
Larvae entirely orange-yellow with areas of red at the bases
of black spines.
Striped Larvae Final Instar
21 °C. and below
Ground colour black broken in each abdominal segment by
stripes of yellow which pass from the mid-dorsal line down to
the laterial ridge. Two of the yellow stripes are narrow and
positioned adjacent to the intersegmental membranes, i.e. they
are positioned at the anterior and posterior of each segment.
The third yellow stripe is broader and positioned slightly an-
terior to the centre of each segment. As in the plain larvae,
these also have yellow patches dorsally situated in the thoracic
segments.
24 °C.
The black areas become slightly tinted with orange especially
the areas adjacent to the mid-dorsal line and lateral ridge.
27-32°C.
The black areas become entirely orange-red in colour.
10
McLEOD
/. Res. Lepid.
Figs. 7-12. Precis octavia Cram.
Upperside, left; underside, right. 7 and 8, f. transiens Wichgraf 9 ; 9 and
10, f. transiens Wiehgraf ; 11 and 12, f. nairohicus (f. nov. ) ^ Paratype.
7 (1): 1-18, 1968
PRECIS OCTAVIA
11
Pigmentation of Pupae
Pupae occurred in four different colour forms which were in
no way related to colour forms of larvae or adults. Also they
did not correspond with any of the environmenal factors or back-
ground colour of their cages. The four colour forms were dark
brown, light brown, mottled and gold.
(Colour Plate 1, Fig. 4)
Pigmentation of Adults
Results obtained are summarised in Tables 2 and 3. These
indicate that neither the light duration nor humidity affect the
pigmentation of the adult. Temperature changes almost exactly
correspond to the differences in pigmentation of the adults. At
the higher temperatures, 27-32 °C. f. natalensis was produced,
and at the lower temperatures, 10-16 °C. f. sesamus was produced
irrespective of the form of the parent or sex of the individual.
A complete range of intermediate forms was bred at tempera-
tures between 18-24° C. A selection of these is illustrated here
in Figures 3-26. Some of these intermediate forms do not cor-
respond with the description of f. transiens (Wichgraf 1918). 1
therefore make descriptive notes here on three new forms as
well as the three forms already described.
Many lepidopterists feel that separate names should not be
given to temperature forms. In this case I feel that because of
the extreme differences within this species, the various forms
should be named and described for ease of reference.
DESCRIPTIVE
1. f. natalensis Staudinger (Figs. 1-2)
Upperside. Ground colour strongly red with a black
margin along the outer border. All discal spots are black and
do not possess pupils. Those of cellules 5 and 6 are larger. No
blue scales are present on the proximal side of the discal spots.
A dark brown area passes from the base of the forewing to half
way along the inner margin and connects with the first trans-
verse bar of the cell but not the second.
Underside. Black areas occur at the bases of all wings. These
black areas contain four orange patches on the hindwings.
Ground colour pinkish red and wings are not demarcated into
halves.
12
McLEOD
/. Res. Lepid.
Figs. 13-18. Precis octavia Cram.
Upperside, left; underside, right. 13 and 14, f. nairobicus (f. nov. ) Holo-
type $ ; 15 and 16, f. susani ( f. nov. ) Paratype 9 ; 17 and 18, f. susani
( f . nov. ) Holotype $ .
7 (1): 1-18, 1968
PRECIS OCTAVIA
13
Note. Some specimens at first sight appear to be of f. natal-
ensis but possess blue or more rarely white pupils to the discal
spots of cellules 5 and 6. It will be found on close examination
with a lens that blue scales are present on the proximal side of
discal spots of cellules 1 and 2 of the forewing. These specimens
are therefore of f. transiens Wichgraf. Generally all specimens
with blue or white pupils to discal spots in cellules 5 and 6 are
intermediates or f. sesamns.
2. f. transiens Wichgraf (Figs. 3-10)
Upperside. Ground colour red with a black margin along
outer border. Small areas of blue scales occur on the proximal
side of the discal spots in cellules 1 and 2 only of the forewing.
Discal spots of cellules 5 and 6 of the forewing with white or
blue pupils. Dark brown areas occur on the distal side of the
second transverse bar of the cell of the forewing and these
may or may not connect up with those of the inner margin.
Underside. Basal half of each wing tends to be demarcated
from the distal half, and is dark brown/ black in colour com-
pared with the pink/ red of the distal half. Marginal band broad
with two rows of blue streaks on upperside and underside.
3. f. nairobicus. f. nov. Holotype S Allotype 9
Bred from wild-caught 9 Karen, Nairobi, Kenya.
Paratypes 9 $ 5 9 , as Holotype and Allotype, in author’s col-
lection.
Upperside. As in f. transiens but blue areas occur on the prox-
imal side of all the discal spots of the forewing and may also
occur on the hindwing. These blue areas remain separate in
each cellule and do not join together. Discal spots in cellules 5
and 6 of the forewing possess white pupils .
Underside, As in f. transiens.
4. f. susani
Holotype S
Allotype 9
Paratypes —
f. nov. (Figs. 15-24)
Bred from wild-caught 9 Karen, Nairobi, Kenya
10 7 9 as Holotype and Allotype, in author’s
collection.
Upperside. Ground colour red. Blue areas are present on the
proximal side of all discal spots of the forewings and hindwing.
In the forewings the blue areas in cellules 1-6 join together to
form a bar. In the hindwings the blue areas tend to be oval and
separate in each cellule, but in more extreme forms the blue
areas unite here also. The areas of dark brown positioned dis-
14
McLEOD
/. Res. Lepid.
Figs. 19-24. Precis octavia Cram.
Upperside, left; underside, right. 19 and 20, f. susani (f. nov. ) Allotype 9 .
21 and 22, f. smani ( f . nov.) Paratype ^ ; 23 and 24, f. msani (f. nov.)
Paratype ^ .
7 (1): 1-18, 1968
PRECIS OCTAVIA
15
tally to the second transverse bar of the cell of the forewing,
join up with those running along the inner margin from the
base.
Underside. Basal half of all wings sharply defined and bound-
ed by a curved dentate line. The distal halves remain pinkish
red in colour but the basal halves are dark brown/ black.
5. f.miotoni f. nov. (Figs. 26-26)
Holotype $ In author’s collection.
Paratype 1 S Both bred from insects wild — caught in Karen,
Nairobi, Kenya. Obviously intermediate between f. sesamus and
f. susani but the areas of blue scales on the proximal side of the
discal spots are diffused amongst the red scales instead of occur-
ring in definite areas. This results in an overall lilac appearance.
The area discal to the discal spots of cellules 1-4 remains red
traversed by the dark veins 2-4. Discal spots in cellules 5 and 6
of the forewings possess white pupils.
6. f. sesamus Trimen (Figs. 27-28)
Upperside. Ground colour blue with a tendency to purple.
Forewings project slightly at the extremity of vein 6. Discal
spots in cellules 1-4 black, in cellules 5-6 white pupilled. Distal
to the submarginal spots of cellules 1-4 are four large red spots.
Marginal band broad with two rows of blue streaks. Basal half
of all wings dark brown.
Underside. Ground colour dark brown/ black with perhaps
traces of pinkish red in cellules 2-3. The basal half of each wing
is bounded by a curved dentate line. Discal spots in cellules
5-6 possess white pupils. Discal spots in cellules 1-4 with or
without white pupils. Note. In extreme forms the discal spot
in cellule 4 of the upperside of the forewing also has a white
pupil (Fig. 29) and lines are more dentate especially those
demarcating the basal from the distal halves of the wings. The
forewings project at the extremity of vein 6 and the ground
colour is more of a definite blue.
SUMMARY & CONCLUSIONS
A brief distribution of Precis octavia sesamus in Africa is
given. Meteorological data is quoted for an area of Kenya in
which both extreme forms of the butterfiy occur. Previous
studies of this species are mentioned.
Experiments are outlined in which numbers of Precis octavia
sesamus were reared under controlled environmental conditions.
The environmental factors controlled were humidity, light dur-
16
McLEOD
/. Res. Lepid.
Figs. 25-30. Precis octavia Cram.
Upperside, left; underside, right. 25 and 26, f . miotoni ( f. nov. ) Holotype
$ ; 27 and 28, f. sesamus Trimen. ^ ; 29 and 30, f. sesamus Trimen 9 .
7 (1); 1-18, 1968
PRECIS OCTAVIA
17
ation and temperature. (Food supply had previously been
shown to be unrelated to pigment changes .
Although numbers of insects were not large, the author con-
siders the evidence sufficient to conclude that:
a) The only environmental factor to affect the pigmentation
of F. ovtavia sesamus is temperature.
b) Temperature, as well as controlling the pigmentation of the
imago, also controls the pigmentation of the final instar larva.
c) The pigmentation of the pupa is unrelated to temperature.
REFERENCES
BUTLER, A. G. 1901. A Revision of the butterflies of the genus Precis.
Ann. Mag. Nat. Hist. 7. p. 205.
CLARK, G. C. and Dickson, C. G. C. 1957. Life History of Precis octavia
(Cram) /. Ent. Soc. S. Afr. Vol. 20, No. 2.
FISCHER, E. 1895. Transmutation der Schmetterlinge in Folge Tempera-
turveranderungen. Berlin.
KETTLEWELL, H. B. D. 1963. The genetical and environmental factors
which affect colour and pattern in Lepidoptera with special reference
to migratory species. Entomologist 96. p. 127.
LECLERCQ, J. 1946. Some effects of atmospheric humidity on two
Nymphalidae. Aglais urticae L. and Araschnia levana L. Proc. R. Ent.
Soc. Lond. 87-88
MARSHALL, G. A. K. and POULTON, E. B. 1902. Trans. Ent. Soc.
Lond. p. 414-460.
MERRIFIELD, F. 1890. Systematic temperature experiments on some
Lepidoptera in all their stages. Trans. Ent. Soc. Lond. p. 131-159.
1891. Conspicuous effects onthe markings and colouring of Lepi-
doptera caused by exposure of the pupae to different temperature
conditions. Trans. Ent. Soc. Lond. p. 155-168.
1893. The effects of temperature in the pupal stage on the colour-
ing of Pieris napi, Vanessa atalanta, Chrysophanus phlaeas and Ephyra
punctaria. Trans. Ent. Soc. Lond. p. 55-67.
1894. Temperature experiments in 1893 with several species of
Vanessa and other Lepidoptera. Trans. Ent. Soc. Lond. p. 425-438.
1906. An address read before the En.t Soc. Lond. 17th Jan. 1906.
“Effects on living things of Temperature.”
1911. Experimental Entomology, Factors in Seasonal Dimorph-
ism. (Premier Congres International d’Entomologie 1910, Bruxelles).
PINHEY, E. C. G., 1949. Butterflies of Rhodesia, p. 89. Salisbury, Rho-
desia.
1965. Butterflies of Southern Africa. Johannesburg, S. Africa.
ROTHSCHILD. W. 1918. Precis octavia variation. Proc. Ent. Soc. Lond, 5,
SEITZ, A. 1925. The Macrolepidoptera of the World. Vol. VIII. The Afri-
can Rhopalocera. Stuttgart.
STANDFUSS, M. 1894. Ueber die Griinde der Variation imd Aberration
des Falterstadiums bei den Schmetterlingen. Leipzig.
1924. Bestimmungsfaktoren des Zeichnungsmusters beim Saison-
Dimorphismus von Araschnia levana prorsa. Biologisches Zentralblatt
44: 173-188.
1900. Synopsis of experiments in hybridisation and temperature
made with Lepidoptera up to the end of 1898. Part 1, Entomologist
33.
1901, Ditto. Part 2, Entomologist 34.
18
McLEOD
/. Res. Lepid.
TRIMEN, R. 1883. Trans. Ent. Soc. Lond. p. 347.
1887. South African Butterflies. A Monograph of the Extra Trop-
ical Species.
WEISMANN, A. 1896. New Experiments on the Seasonal Dimorphism of
Lepidoptera. Translation W. E. Nicholson. Entomologist.
1875. Ueber den Saison-Dimorphismus der Schmetterlinge. Leip-
zig.
1880. Studies in the theories of descent. Part 1. On Seasonal
Dimorphism of Butterflies. Translated by R. Meldola.
WICHGRAF, E. 1918. Neue Afrikanische Lepidopteren. Int. Ent. veit-
schr. 12.
ACKNOWLEDGEMENTS
The author is indebted and grateful to the following for per-
mission to carry out experiments in their laboratories. Mrs.
E. C. van Someren, Division of Insect Borne Diseases, Medical
Department, Nairobi, Kenya; Mr. G. R. Cunningham - van Som-
eren, East African Research Unit, Karen, Nairobi, Kenya; Dr.
M. Coe, Zoology Department, University College, Nairobi, Ken-
ya; Special thanks also to Dr. V. G. L. van Someren, ''the Sanc-
tuary,” Karen, for his encouragement and helpful advice.
EDITOR’S NOTE. It is planned at some time in the near fntvire to illustrate the larvae,
pupae and adidt forms in color.
Journal of Research on the Lepidoptera
7(1): 19^22, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 196H
ECOLOGICAL AND DISTRIBUTIONAL NOTES
ON EREBIA DISA (SATYRIDAE)
IN CENTRAL CANADA
JOHN H. MASTERS*
Research Associate, Carnegie Museum., Pittsburgh, Pa.
Erebia disa (Thunberg) is a circumpolar species with several
geographic subspecies including mancinus Doubleday which
occurs in central Canada west to the Rockies. As late as 1936
(Warren), E. disa was not known to occur east of the Alberta
Rockies in North America; however, Brooks (1942) recorded it
from Aweme and Gillam., Manitoba and Riotte (1959) recorded
it from five localities in extreme northern Ontario east to Smoky
Falls. More recently its range has been found to be more ex-
tensive and complete. Riotte (1962) recorded it at Hymers,
Ontario, near the Minnesota border; John Polusny and C. S.
Quelch {in lift.) captured specimens in Southwest Manitoba at
Sandilands Provinical Forest during June 1967; I captured three
specimens at Riding Mountain National Park, Manitoba on 25
June 1967; and during June and July of 1968, Patrick J. Conway
and I filled in many gaps in the range by discovering eight col-
onies in Manitoba and Ontario (figure 1.),
I found Erebia disa restricted to black spruce/sphagnum bogs
and especially those bogs having tall, dense stands of pure spruce.
In this habitat, disa was encountered among the larger spruce,
but a few strays were observed in more open bog areas or along
roads bordering bogs. Ehrlich (1956) found that E. disa in Alas-
ka always appeared to be associated with spruce forest, but
noted strays in sedge marshes or crossing roads. Ehrlich also
noted numbers of disa sucking moisture from a damp road at
mile 1316 on the Alaska Highway. In the Palearctic Region,
E. disa is usually depicted as being associated with marshes,
however, in Norway, Sheldon (1913) found that disa preferred
a wet “moor” overgrown with vaccinium rather than nearby
swamps and marshes.
’ Home address: P. O. Box 7511, Saint Paul, Minn.
19
20
MASTERS
/. Res. Lepid.
Figure 1. Central Canada showing expected range of Erehia disa man-
cinus Doubleday ( shaded area ) , perviously known localities ( circles )
and newly reported localities ( triangles ) .
7 (1): 19-22, 1968
EREBIA DISA
21
Erehia dim has a decided tendency to shun bright sunlight
and, for the most part, will fly in the morning before 11:00 A.M.
and again in the evening after 4:00 P.M. On cloudy days they
will fly through the noon hours, but on sunny days can be
flushed with difficulty from foliage at the base of spruce trees.
E. disa flies slow and. steady about three feet off of the ground
and perfers lighting in partial sunlight on low foliage at the base
of spruce trees. The slow flight can be deceptive as unlike
Oeneis jutta (Hubeer), which darts back and forth in rapid
flight, disa will maintain a linear direction through a bog and
can quickly outdistance a collector. Because Erehia dha is a
forest dweller, I thought that it might exhibit some degree of
territorialism, as was noted in forest dwelling Oeneis (Masters
and Sorensen, 1969 ) , but territorial behavior was ' not detected.
In Satyridae, territorialism and “hilltopping” seem to be closely
related; Shields ( 1968 ) includes Erehia among a list of genera
that “are apparently devoid of hilltopping species.”
Oeneis jutta was the only other species that always seemed
to be associated with Erehia disa (also noted by Ehrlich, 1956),
Erehia disa might have a biennial flight as do several Palearc-
tic Erehia including E. claudina (Bkh.) and £. Ugea (L. ). The
late Richard J. Fitch, formerly of Rivercourse, Saskatchewan,
first collected E. disa in spruce bogs near Harlan, Saskatchewan
in 1942 and thereafter only seemed to^ encounter it in even num-
bered years (Masters, 1968). While E. disa has now been taken
in consecutive years in the same area in Manitoba-— it has not
,i)een retaken in the same bogs. In fact, I found it wanting in
1968 in the same Riding Mountain bog where I had found it in
1967 and John Pokisny (in Utt.) was unable to retake it in 1968
in the Sandilands bog where he found it the previous year. The
pattern with biennial Erehia in Europe has nearby ’ colonies ran-
domly alternating with each other on the year of flight and it
appears that his might be the case with Erehia disa mancinus.
This is quite different from the pattern in biennial appearing
Oeneis where populations over extensive areas are on the same
cycle and alternation occurs only across a natural barrier such
as a mountain range or desert.
Erehia disa has not yet been recorded in the United States
(exclusive of Alaska). It almost certainly occurs in the norther
tier of counties in Minnesota. The presence of disa at Aweme,
Manitoba suggests that it might also occur in the Turtle Moun-
tain area of North Dakota and there is also a good possibility
22
MASTERS
/. Res. Lepid.
that it will be found in the Rockies of western Montana. The
map ( figure 1 ) shows the known localities and suggested range
of Erebia disa in the central area. The Alberta localities are
from Bowman ( 1951 ) ; the data for the other localities follows :
MANITOBA: Gillam (G. S. Brooks, 1942 and F. H. Chermock,
1967); Thompson (F. H. Chermock, 1967); The Pas, 1 July
1968, J. H. Masters and P. J. Conway; Harte Mountain, Porcu-
pine Prov. Forest, 2 July 1968, J. H. Masters and P. J. Conway;
Favel River, Duck Mtn. Forest Reserve, 3 July 1968, J. H. Mas-
ters and P. J. Conway; Blue Lakes, Duck Mtn. Prov. Park, 5
July 1968, J. H. Masters and P. J. Conway; Lake Jane, Riding
Mtn. National Park, 25 June 1967, J. H. Masters; Aweme, N.
Criddle (Brooks, 1942); Sandilands Provincial Forest (2 loca-
tions), June 1967, C. S. Quelch and John Polusny; Whiteshell
Provincial Park, 29 June 1968. J. H. Master's.
ONTARIO: Reed Narrows, 29 June 1968, J. H, Masters; Long-
bow Corners, 29 June 1968, J. H. Masters and P. J. Conway;
Hymers (Riotte, 1962); Favourable Lake (Riott^ 1959); Gerald-
ton (Riotte, 1959);, Nakina (Riotte, 1959); Ogoki Post (Riotte,
1959); Smoky Falls (Riotte, 1959).
SASKATCHEWAN: Harlan, June, R. J. Fitch (Masters, 1968);
North shore of of North Saskatchewan River, 20 miles north of
Lloydminster, June, R, J. Fitch (Masters, 1968); 5 miles east of
junction of highways 165 and 106, 10 June 1968, J. S. Nordin.
REFERENCES CITED
BOWMAN. K., 1951, An annotated list of the Lepidoptera of Alberta.
Canadian Journal of Zoology 29: 121-165.
BROOKS, G. S., 1942. A check list of the butterflies of Manitoba. Canadian
Ent. 74; 31-36.
CHERMOCK, P. W. and F. H. CHERMOCK, 1968. Churchill. Bull As-
soc. Minnesota Ent. (Minneapolis) 2: 33-39.
Satyridae) in Northwestern America. Ent. News 67: 29-36.
MASTERS, J. H., 1968. R. J. Fitch’s list of Saskatchewan butterflies. The
Blue Jay ( Regina Saskatchewan ) 26: 176-181.
MASTERS, J. H. and J. S. SORENSEN, 1969. Field observations on forest
Oeneis ( Satyridate). J. Lep. Soc. 23: in press.
RIOTTE, J. C. E., 1959. Revision of C. J. S. Bethime’s list of the butter-
flies of the eastern provinces of Canada as far as northern Ontario is
concerned. Ontario Field Biol. 13: 1-18.
RIOTTE, J. C. E., 1962. First additions to the northern Ontario list of
butterflies. /. Lep. Soc 16: 243-245.
SHELDON, W. G., 1913. The Lepidoptera of the Norwegian provinces
of Odalen and Finmark. The Entomologist 46: 11-15.
SHIELDS, O,, 1968. An ecological study of summit congregation behavior
of butterflies on a southern California hill, /. Research Lepidoptera
6: 69-178.
WARREN, B. C. S., 1936. Monograph of the genus Erebia. The Oxford U.
Press (London) pp. 1-407, 104 pi.
Journal of Research on the Lepidoptera
7(1): 23-28, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
METHODS FOR STUDYING THE
CHROMOSOMES OF LEPIDOPTERA
THOMAS C.,EMMEL
Department of Zoology, University of Florida, Gainesville 32601
Chromosome information has been published on over two
hundred butterfly species in the Palearctic region (de Lesse,
1960, and included references). A smattering of chromosome
counts has been published on African species of Rhopalocera
(de Lesse and Condamin, 1965, 1966), and an initial survey of
105 Nearctic and northern Neotropical species has been given by
Maeki and Remington ( 1959, 1960a, 1960b, 1960c ) . Some Aus-
tralian species have been studied cytologically (Emmel & Mc-
Farland, unpublished). Recently, de Lesse (1967) published a
list of the chromosome numbers of 284 Neotropical Rhopalocera,
all from South America; these include 14 species already counted
by Maeki and Remington from northern Mexico. These refer-
ences are the only major publications to date on butterfly chrom-
osomes. Yet with the exception of Drosophilidae no other large
group of animals approaches the degree of cytotaxonomic knowl-
edge (over 700 species) we now have of the Rhopalocera
(Maeki and Remington, 1960c). The karyotypes of the moths
are almost totally unknown.
In the course of extensive investigation of the karyotypes of
Neotropical and Nearctic butterfly species, the author has devel-
oped a simplified set of techniques for obtaining and studying
the chromosomes of Lepidoptera. The purpose of the present
paper is to outline these methods.
COLLECTION AND PRESERVATION OF TESTES
Chromosomes are most easily studied in dividing cells in the
testes of male butterflies. Meiosis usually continues there for
some time after eclosion (up to several months in Heliconius
species ) , the haploid number is easier to observe than the diploid
complement of mitotic somatic cells, and meiosis in the eggs of
a female only occurs singly during the short interval of sperm
combination with each egg (see also Maeki and Remington,
1959).
23
24
T. C. EMMEL
/. Res. Lepid.
In almost all butterflies, the two testes are fused laterally and
located at the top of the abdomen, beneath the junction of the
third and fourth abdominal segments counting forward from the
genitalia (the easiest way to count in the field). In the large
sulfur genus Phoebis and certain other Neotropical butterflies,
the testes are placed in the top of the clasping apparatus ( term-
inal abdominal segment).
Location: The roundish or oblong testes are always in the
center and have two long tubes attached to their joined
base.
Colon They are usually rose or red, but may be greenish,
black, yellow, or even clear in many lycaenids, satyrids,
and certain Papilios.
Size: This varies greatly, depending on age (decreasing
size in older individuals ) and the species. Heliconius and
danaids have testes up to 2 mm in diameter. In some
satyrids and lycaenids (blues), they may be only 0. 1-0.2
mm in diameter.
Directions for Removal of Testes:
To remove the testes, hold the male butterfly in the left hand
with its wings above the thorax; shove the abdomen at an angle
downwards with a free finger. With fine watchmaker’s forceps
(No. 5 size is best) in the right hand, tear open a slit in the top
of the abdomen at the junction of the third and fourth segments
back from the claspers (Fig. 1). The distinctively-colored testes
should pop into view immediately; the other abdominal contents
are yellowish, white or translucent.
Pull off the testes and place into a screw-cap vial (one- or
two-dram size are satisfactory) containing a 3:1 mixture by
volume of absolute ethyl alcohol and glacial acetic acid (freshly
mixed or not more than a few hours old). Push the abdominal
contents back in the male’s abdomen, close the slit, and store
the specimen in an envelope for future reference. Put a label
with a pencil- written code number in the vial with the testes;
put the same number in ink on the glassine envelope containing
that male specimen; and enter the same number and collection
data in a permanent notebook. The wing condition or apparent
age of the male can be noted, also. Use a different vial for the
testes of each individual butterfly.
The testes can be stored in the original vials until chromosome
squashes are made; it is not necessary to transfer them to alcohol
for storage. The vials should be refrigerated and if possible
7 (1): 23-28, 1968
CHROMOSOMES
25
stored at freezing temperatures at the earliest opportunity. The
testes will give satisfactory chromosome perparations even after
two years of frozen storage, though faster processing is recom-
mended.
Because some individuals in a population may not be under-
going active spermatogenesis at the same time as others, it is
advisable to collect 5-10 testes for each species ( or more if there
is known to be a variable number of chromosomes in popula-
tions of that species).
The collection of testes from live males does not have to be
done immediately upon netting of the specimens. Males can be
left alive (unpinched) in glassine envelopes all day as long as
they do not get overheated, and the testes can be removed in
the evening as each male is killed.
26
T. C. EMMEL
]. Res. Lepid.
SQUASH TECHNIQUE FOR CHROMOSOME STUDIES
The following squash technique is the simplest and fastest one
to use to obtain good preparations of chromosomes which then
may be photographed and drawn via camera lucida for perman-
ent record. This procedure does not produce permanent slides,
though these squash preparations may be sealed with clear nail
polish and held at freezing temperatures for several years if
desired. However, slides may be made permanent by dehydrat-
ing the preparations in a series of alcohol concentrations (Guth-
rie, Dollinger, & Stetson, 1965).
1. After removing testes from fixative in the vial or from a
freshly-killed male, place on a clean slide in a drop of Lacto
Orcein Stain (see Appendix I).
2. Macerate the testes with watchmaker’s forceps. Allow to
Fig, 2. General field of dividing testicular cells at 400x microscope mag-
nification. Eurenia sp. (Pieridae), San Vito de Java, Puntarenas
Province, Costa Rica (n=:29). It is relatively rare to obtain cells
with all chromosomes in the same plane of focus; only one or two of
the sets of chromosomes in this field of view could probably be counted
accurately (at higher magnifications).
Fig, 3, Highly magnified haploid set of chromosomes (n=:21) from a
testis cell of a Heliconius melpomene male ( Nymphalidae: Heliconi-
inae), Osa Peninsula, Puntarenas Province, Costa Rica. The line of
chromosomes at left represents an equatorial view of chromosomes
on a metaphase spindle in a neighboring cell. The microscope magni-
fication for this photograph was lOOOx.
7(1): 23-28, 1968
CHROMOSOMES
27
stand for at least 5 minutes. (A longer exposure to the stain,
up to several hours, gives better results. Cover stain drop
with a small watch glass to prevent evaporation).
3. Place cover slip over drop of stain. Tap the top of the cover
slip to spread out cells.
4. Put a paper towel or filter paper on top of cover slip and
squash by thumb pressure over cover slip. ( Or, the slide may
be inverted, placed on a sheet of glass with paper toweling
above and below, and thumb pressure applied to squash.)
A Carver Laboratory Press may be used to insure a uniform,
well spread chromosome preparation.
5. Remove excess stain around edges of cover slip with filter
paper and examine slide under low power (lOOx) to locate
areas of dividing cells.
Dividing cells may be examined under oil-immersion for
counting, description and photography. The author uses a Carl
Zeiss Research Microscope STANDARD WL fitted with
plan-apochromatic flat-field objectives and automatic camera.
An oil-immersion Planapo lOOx objective is used for critical
observation and photography. Total magnifications of at least
lOOOx are needed for studying the tiny chromosomes of the
Lepidoptera.
Examples of the appearance of areas of dividing testicular
cells and of chromosomes at high magnification are given in
Figures 2 and 3.
ACKNOWLEDGMENTS
This research has been supported by the N.I.H. Genetics
Training Grant to the University of Texas at Austin (1968-1969)
and by N.S.F. Grant GB8442. I thank Dr. Guy L. Bush for his
great assistance at the University of Texas in the initiation of
this chromosome research on Lepidoptera and for his continued
interest and support.
28
T. C. EMMEL
J. Res. Lepid.
LITERATURE CITED
DE LESSE, H. 1960. Speciation et Variation chromosomique chez les
Lepidopteres Rhopaloceres. Annales des Sciences Naturelles, Zoologie,
12e Serie, 2(1): 1-123.
DE LESSE, H. 1967. Les Nombres des Chromosomes chez le Lepidopteres
Rhopaloceres Neotropicanx. Ann. Soc. ent. France (N.S.), 3(1): 67-
136.
DE LESSE, H., and M. CONDAMIN. 1965. Formiiles chromosomiques de
quelqiies Lepidopteres Rhopaloceres dii Senegal et de Cote d’Ivoire.
Bulletin I.F.A.N., 27, A(3): 1089-1094.
DE LESSE, H., and M. CONDAMIN. 1966. Formules chromosomiques de
quelques Lepidopteres Rhopaloceres d’Afrique Centrale. Ann. Soc. ent.
France (N.S.), 11(2): 349-353.
GUTHRIE, W. D., E. J. DOLLIINGER, and J. F. STETSON. 1965.
Chromosome studies of the European corn borer, Smartweed borer,
and Lotus borer (Pyralidae). Annals Ent. Soc. Arner., 58(1): 100-105.
MAEKI, KODO, and CHARLES L. REMINGTON. 1959. Studies of the
chromosomes of the North American Rhopalocera. 1. Papilionidae.
Journ. Lepid. Soc., 13: 193-203.
. 1960a. Idem. 2. Hesperiidae, Megathymidae, and Pieridae. Journ.
Lepid. Soc. 14: 37-57.
. 1960b. Idem. 3. Lycaenidae, Danaeinae, Satyrinae, Morphinae.
Journ. Lep. Soc., 14: 127-147.
. 1960c. Idem. 4, Nymphalinae, Charaxidinae, Libvtheinae. Journ.
Lepid. Soc., 14: 179-201.
APPENDIX I
iMcto Orcein Stain
I gm Orcein
40 cc Glacial Acetic Acid
10 cc Lactic Acid (undiluted)
50 cc Water (distilled)
firing to a boil, let mixture stand in flask overnight, then filter
with Whatman No. I filter paper to remove crystals. Store stain
in small nose-dropper bottles.
Journal of Research on the Lepidcptera
7(1): 29^30, 1968
1160 W. Orange Grove Ave., Arcadia, California, US. A. 91006
@ Copyright 1968
NOTE ON VITAL STAINING
OF ACTIAS LUNA SILK
JOHN M. KOLYER
55 Chimney Ridge Drive, Convent, New Jersey 07961, U’.S.A.
Vital staining results with Pieris rapae (Linnaeus) and
CoUas species have been reported (Kolyer, 1965 and 1966). In
P. rapae, the dye neutral red imparted a red tint to all stages,
and Sudan black B gave bluish larvae, greenish-black pupae,
and blue internal color in adults. Silkworms fed neutral red spin
“bright red” cocoons ( Edwards, 1921 ) .
In the present work, the dyes fed to Actias luna (Linnaeus)
larvae (5th instar) were neutral red, Colour Index No. 50040,
total dye content 88%, and Sudan black B, Colour Index No.
26150, both obtained from Allied Chemical Corp., New York
City. These were ground and blended with P-12 Davenite mica
(325 mesh; Hayden Mica Co., Wilmington, Mass.) at 3 parts dye
per 97 parts mica, and the blend was rubbed on the underside
of hickory leaves at about 18 mg. blend/inc surface. Incident-
ally, with stems in water the leaves kept well below 80 °F at
about 50% rek humidity but wilted rapidly at 85-90 °F.
Neutral red showed toxicity and caused pronounced inhibition
of growth, as noted for the butterfly species. While no mortality
occurred among control larvae, of 16 larvae fed the dye for 2-8
days (followed by feeding undyed leaves if larvae hadn't died)
only 2 survived to produce cocoons; these were fed dye for 2
days, at which point one began spinning while the other began
a day after being transferred to undyed leaves. Success seemed
to depend on feeding dye only long enough to saturate the body
with the color ( 2 days, or even as little as 12 hours with voracious
feeding) and on choosing larvae almost ready to pupate.
29
30
KOLYER
J. Res. Lepid.
Four larvae were fed Sudan black B for 7 days and then
transferred to undyed leaves. Only one survived and spun a
cocoon. In this case, desorption of dye from the body was indi-
cated; feeding on undyed leaves caused gradual loss of the deep-
er green shade given by the dye. Similar reversibility of neutral
red has been noted for the wild silkworm Attacus Orizaba (Ed-
wards, 1921 ) as well as in the cited work with butterflies. The
cocoon was uncolored like that of a control.
The two strongly-pink or rose colored cocoons from the neu-
tral red experiment were opened to disclose dead larvae. After
discarding the latter and picking all leaf fragments from the silk,
the cocoons weighed 70 and 79 mg. Each cocoon was assayed
for neutral red by triturating a sample (37-39 mg.) with 6 ml.
of concentrated (37-38%) aqueous HCl, filtering, and measuring
optical density at 725 millimicrons with r. Bausch and Lomb
Spectronic 20 Colorimeter. A calibration curve was constructed
using known concentrations of dye in the HCl solution. The
result was 1,3 ±: 0.1% neutral red (as “total dye”^ in the cocoons.
Experiments with Attacus Orizaba ( Edwards, 1921 ) have shown
that the neutral red in the cocoon indeed is transferred to the
sericin through the insect’s body rather than being picked up
externally during spinning.
LITERATUBE CITED
EDWARDS, W. F., 1921. Feedin^^ dvestiiffs to silkworms. Textile World,
60: 1111-1113.
KOLYER, J. M., 1965. The feeding of coloring matters to Pieris rapae
larvae. Jour. Res. Lepid., 4(3): 159-172.
1966. Vital Staining of Colins philodice and C. euriitheme. Jour.
Res. Lepid., 5(3): 137-152.
Journal of Research on the Lepidoptera
7 (l):31-34, 1969
1160 W. Orange Grove Ave., Arcadia, California, US. A. 91006
© Copyright 1968
PRESENT AND ICE AGE LIFE ZONES
AND DISTRIBUTIONS
WILLIAM HOVANITZ
In describing the distributional ranges of any insect, it is
very desirable that some means of rapid correlation with a
climatic zone (or vegetational zone) be available. Many years
ago, Merriam (1898) found a scheme of “life-zones” useful for
the purpose of describing the distribution of North American
mammals; such zones were heavily relied on by other vertebrate
zoologists as an aid toward the ecological description of distri-
bution patterns (see Grinnell and Storer, 1924 and Hall and
Grinnell, 1919).
In addition to existing life zones, some idea of the distribu-
tion of the zones in the immediate geological past is desirable.
L. S. Dillon (1956) has attempted to show the relationship
between the life zones of the present and of the ice age. The
reader should refer to his paper for an excellent discussion of
the subject. Nevertheless, since that paper is not radily available
to many readers of this Journal, two significant maps from that
paper are here redrawn and reproduced in color. The first is a
simplified map of Merriam’s Life Zones of the present and the
second is a map showing the presumed or hypothetical location
of the same life zones during the Wisconsin ice age.
Of prime importance to formulation of conclusions on the
possible reasons for some idiosyncrasies in present day distri-
butions, one major fact emerges. Gontinuity of distribution be-
tween Alaska and the more southerly parts of North America
which existed before and now exist after the ice ages were com-
pletely broken for a long period of time. For example, if one
took a group of Golias such as Vaccinium feeders which are near-
ly circumpolar, it is apparent that these could extend without
much discontinuity from Europe across Asia into Alaska and
across northern North America. The ice barrier of the Pleistocene,
however, effectively isolate the Alaska end of the Asian distri-
bution from the more southerly parts of the North American dis-
tribution and the possibility of local races or species developing
31
32
W. HOVANITZ
J. Res. Lepid.
FIG. 1 Present day life zones (simplified from Merriam and Dillon). Purple: Arctic-
alpine (tundra); blue: Hudsonian; green: Canadian; yellow: Transition; red-orange:
Upper Austral (Sonoran); red-magenta: Lower Austral (Sonoran). Ice not shown in
Cireenland and adjacent islands.
7 (l):31-34, 1969
DISTRIBUTIONS
33
FIG. 2. Hypothetical life zones of the Wisconsin ice age. Same legend as Fig. 1
except white in north designates arctic glaciation. Both these figures have been redrawn
from Figures 10 and 11 of Dillon (1956) and reproduced in color by permission of
Lawrence S. Dillon and Science, published by the American Association for the Ad-
vancement of Science, Washington, D.C.
34
W. HOVANITZ
/. Res. Lepid.
would be strong. Colias paleano (The Eur-Asian Vaccinium
feeder ) would remain in that sector but new forms would
develop through genetic isolation and habitat selection south-
east of the ice. For example, Colias interior might have devel-
oped spanning the North American continent, or Colias minisni
in the southern Canadian Rockies, or Colias behri in the Sierra
Nevada, or Colias pedidne in Labrador and possibly Colias
scudderi in the southern Canadian Rockies. Movement north-
ward of Colias interior and southward of Colias palaeno
after the retreat of the ice would create a zone of hybridization
if they retained the same ecological niches, and were still inter-
fertile. But such was not the case and from the mouth of the
MacKenzie River to Manitoba, they overlap with no known
mixing. Two other species (Colias hecla and C. nastes) however
have been confused by the new habitats following retreat of
the glaciers and have partly blended together with the forma-
tion of a partial new species (Colias hoofhi) (Hovanitz 1950,
1951 and 1963).
REFERENCES
GRINNELL, JOSEPH and T. T. STOKER. 1924. Animal Life in the
Yoseinite, Univ. Calif. 752 pp.
MERRIAM, C. HART. 1898. Life Zone.s and Crop Zone.s of the United
States. U. S. Dept. Atiric., Div. Biol. Sur. Bull No. 10. 79 pp.
DILLON, L. S. 1956. Wiseonsin elimate and Life Zones in North Ameriea.
Science 123( 3188 ): 167-176.
HOVANITZ, WILLIAM. 1950. The Biology of Colias Butterflies. 1. The
distribution of the North Amereian .Speeies. Wasmann Journal of
Biology 8:49-75.
1951. H. Variation of adult flight in the Aretie and Siibaretie.
Wasmann J. Biol. 9:1-9.
1963. The origin of a sympatrie speeies in Colias through the
aid of natural hybridization. }. Res. Lepid. 1:261-274; 2:205-223.
Journal of Research on the Lepidoptera
7 (1): 35-49, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
Copyright 1968
TRIALS OF SEVERAL DENSITY ESTIMATORS
ON A BUTTERFLY POPULATION
WILLIAM R. HANSON and WILLIAM HOVANITZ
Department of Zoology, California State College, Los Angeles
INTRODUCTION
The density of an animal population is notoriously difficult
to estimate, and new methods are consequently being developed
continually. Since some newer procedures have been tried little
in the field, our objective was to compare several of them to
several older ones.
Some of the extensive literature on population estimation has
been reviewed by Hanson (1967), Southwood (1966), and
Ricker (1958), making further discussion of the theory not now
warranted. The book by Southwood emphasizes entomological
applications, and especially how to obtain reliable data. It
bears repeating that workers have found it considerably easier
to develop the mathematical bases of the estimating techniques
than they have to solve the biological and economic problems
of getting unbiased data in adequate amounts for use in the
estimators.
To compare the estimating procedures, we required to study:
(1) a natural population, (2) one that was fairly dense, (3) a
relatively isolated population, to reduce egress of marked ani-
mals, (4) yet one comprising a highly mobile species, and finally,
(5) a population that could be found close at hand and ap-
proached and captured with a minimum of problems. For these
purposes, the common alfalfa butterfly (Colias eurytheme)
turned out to be very good. The habitat and behavior of the
alfalfa butterfly, among other matters, were discussed by Hova-
nitz (1948).
A suitable population of the butterflies was found in a field
of alfalfa (Medicago sativa) located on an experimental farm
of California State College at Pomona. The field contained 14.2
acres, was rectangular in shape, and was surrounded by grass.
35
36
HANSON-HOVANITZ
J. Res. Lepid.
fallow land, and an orange grove. It appeared to be well iso-
lated from other areas providing habitat for Colias eurytheme.
The alfalfa was somewhat thinly planted and averaged about
12-14 inches tall. The field data were collected on three con-
secutive days, August 13 through 15, 1964. (Further work was
attempted in another alfalfa field in August of 1966 but inade-
quate isolation of the population precluded any reliance on
marking methods. )
METHODS OF GETTING DATA
Throughout the field, two or more workers moved about at
random, netting the butterflies that came within reach. Upon
capture, each butterfly was marked with a spot of nail polish
on the ventral, distal surface of the wing; the butterfly was held
for a few moments to allow the paint to dry and then was
released, in the manner to be described in a forthcoming paper
by Hovanitz. By using several dots, it was easily possible to
show how many times a given individual butterfly had been
captured. Marking was not continued beyond the second day.
Goncurrently with this effort, in a second “experiment,” two
other workers attempted to make total counts on sample plots in
the Cal Poly field. Before our work began, the alfalfa field had
been divided lengthwise into 10 strips, each about 93 feet wide,
by low dikes erected to keep irrigation water in place. As the
observers moved lengthwise along each resulting strip, they
walked 20 long steps ( about 60 feet ) and counted all butterflies
within the resulting “plots,” then stopped and recorded the
insects seen, and continued to repeat this process. The size of
the plots (60 X 93 feet) was determined partly by the fact that
the observers concluded not to count any butterflies that were
more than 60 feet beyond them.
In another experiment, a series of cursory, incomplete counts
was made in this alfalfa field by one observer. In this case the
observer walked rapidly back and forth across the field from
one side to the other. The beginning point and ending point
of each walk were guided on a stake previously set at the middle
of each side. When he crossed the field, the observer’s eyes were
fixed straight ahead on the stake located at the far side, but all
Colias that could be seen within the arc encompassed by the
observer’s vision as he looked straight ahead were included in
the counts. Since the field was crossed 35 times, 35 superfieial
samples of butterflies were gathered.
7 (1): 35-49, 1968
POPULATION ESTIMATORS
37
FREQUENCY OF CAPTURE
The repeated capturing, marking, and releasing of the butter-
flies produced a frequency distribution in which fi butterflies
were caught x times, fs were caught 2x times, and so on up
through f i animals taken x , times for each of the two days on
the Cal Poly field, as is shown in Table 1. (The estimated
abundances of butterflies according to this and all other methods
is shown in Table 2. ) The resulting data were used to estimate
the frequency of the animals seen zero times, i. e., to estimate
the missing class fo in the truncated distribution. After the
number of unseen animals was estimated, obviously it could be
added to the number of those actually seen to give the estimated
total number of butterflies in the whole population.
As was discussed in the earlier review (Hanson, 1967), several
papers give promising procedures for estimating the total abund-
ance, K, of the population from such frequency of capture data.
Among these, the paper by Craig (1953) contained a refined
version of a moment model using data obtained by Hovanitz
(Method 2 of Craig’s paper), which required the data to have
an underlying Poisson distribution; this model was tried on the
data shown in Table 1. A paper by Edwards and Eberhardt
( 1967 ) contained several estimating procedures, among which
was the maximum-likelihood model requiring data coming from
a geometric distribution.
A further procedure mentioned by Edwards and Eberhardt
involved plotting of the capture frequencies on semi-log paper.
It is well known that when a regression relationship is curvi-
linear, it can often be transformed into a linear one by plotting
the logarithm of one or both variables ( see, for example, Bailey,
1959:94). In the familiar expression for the linear regression line
Y z= a -f b X,
Y is the dependent variable; X is the independent variable; a is
the height on the Y axis where the line began, and b is the slope
of the line. One can plot the number of animals captured once
against the number 1, the number captured twice against the
number 2, etc. When semilog paper is used and the number of
animals is plotted on the logarithmic scale (i.e., on the Y axis)
and the number of times that they were captured is plotted on
the equal-interval scale (i.e., on the X axis) a straight line may
result. If the points (X,Y) result in a straight line, then the
transformed statement of the regression equation must have
finallv ended up with the form (see Steel and Torrie, 1960:334):
log Y = a + bX.
38
HANSON-HOVANITZ
/. Res. Lepid.
TABLE 1
The Number of Times That Butterflies (Colias eurytheme) Were Captured
and Marked in an Alfalfa Field Near Pomona, California
X
(Number of Captures
Per Individual)
f
(F requency)
xf
x^f
I. ALL SEXES:
0
--
1
81
81
81
2
35
70
140
3
11
33
99
4
1
4
16
Sum:
128
188
336
n. FEMALES ONLY:
0
---
...
--
1
46
46
46
2
24
48
96
3
10
30
90
4
1
4
16
— —
•
Sum: - - -
81
128
248
7(1): 35-49, 1968
POPULATION ESTIMATORS
39
However, the last equation differs from the expression
used by Edwards and Eberhardt (1967:92), which they got
from the geometric-distribution model, since their expression
had the form
log Y = a 4- X log b .
Therefore, it seems doubtful that plotting the capture frequen-
cies on semilog paper, and the corresponding number of times
each animal was captured on the equal-interval scale, will pre-
serve the meaning of the geometric expression, but statisticians
should investigate the matter further. In any case, the plotting
of our data on semilog paper gave good estimates, as will be
shown later (Figure 1).
Poisson Estimator
The data on captures of Colias eurytheme for use in the
Poisson estimator shown as Method 2 in Craig, 1953) are given
in Table 1, and the estimated number obtained by all methods
are summarized in Table 3. Based on the data for all sexes, the
result for Method 2 gave
K := 1882 / (336- 188) 239.
When the error is expressed as a decimal fraction of the esti-
mated mean according to Craig’s formula the result at the 95%
confidence level is:
Standard Error =<7^/^— 2 ( 239) / 1882 = .1162.
Therefore, the confidence limits became (Table 3)
185 < 239 < 293 .
When only the data for females were used, the estimated
number of females was 137 and its 95% confidence limits were
102 to 172. Because the sex ratio among butterflies seems usually
to be approximately unity, these numbers can be doubled to
give K = 274 and a confidence interval extending from 204 to
344 (Table 3).
As work progressed, some marked butterflies moved out of
the field, and the population started declining. We suspected
that the insects moving out were, as usual, mainly males, which
made the results based on females better than that based on
both sexes combined. Although the real number of butterflies
inhabiting the field when work began was obviously unknown,
the results show that it was approximately 275, and the 95%
confidence limits extended from 200 to 350.
Evidently the data did come from a Poisson distribution or
from one that approximated it tolerably well. The procedure
required that jy not change much from trial to trial, and evidently
40
HANSON-HOVANITZ
/. Res. Lepid.
TABLE 2
The Number of Time s That Free-Ranging Butterflies (Colias eurytheme)
Were Observed on Sample Plots in an Alfalfa Field Near Pomona^ California
X
(Number of Colias
Seen Per Plot)
f
(F requency of
Plots)
fx
(Total Colias)
fx^
0
60
0
0
1
27
27
27
2
16
32
64
3
4
12
36
4
4
16
6^
Sum:
Variance = = 1. 116
111
87
191
Mean = x = . 784
7(1): 35-49, 196H
POPULATION ESTIMATORS
41
this condition was met. The labor of capturing the butterflies
during the hot weather was considerable, yet was small com^
pared to that required to catch and mark animals such as fishes,
birds, and mammals with nets, baited traps, or comparable
means.
In summary, the frequency-obcapture method using the
Poisson model gave good results. Movement of marked butter-
flies off the study area was a problem, just as it is for mark-and-
recapture models or removal models (see Ricker, 1958:86 for
further discussion), making it necessary to work quickly and
to stop as soon as a few of the animals have been captured as
many as four times.
Geometric Estimator
When the same basic data (Table 1) were used in the equa-
tion of Edwards and Eberhardt ( 1967 ) , the results were
128
K = =401
1 ^ (128 --- 188)
Confidence limits were not calculated since no procedure for this
was given by Edwards and Eberhardt.
The estimate of 401 butterflies obtained from the geometric
model was well above what we believed to be approximately
the correct upper bound of 350. Why this model did not give as
good an estimate as the Poisson was not clear, but possibly it
was because the geometric model is more suitable for contagious
(clumped) spatial distributions. However, these butterflies flew
about in an apparently random manner, and gave no evidence
of significant aggregation.
Regression Estimator
When the frequencies-of-capture were plotted on semi-log
paper, the resulting points fell remarkably close to a straight
line (Figure 1). For both sexes combined, the fit was very good
except for the class of four captures per individual, where the
sample size was, of course, very small. When this point was
ignored, the plotted line indicated that the zero class of frequen-
cies was about 162, and that the total population was thus about
290 (Table 3). For females only, the fit of the line was even
better (Figure 1), and it indicated that about 234 animals were
not captured, making the total population about 315 (Table 2).
Both estimates are near what was believed to be the true num-
ber, 275.
Since the method showed promise and could be applied
quickly, it should be tested considerably more. Getting the
42
HANSON-HOVANITZ
J. Res. Lepid.
TABLE 3
Summary of Estimates of the Number of Butterflies (Colias eurytheme )
Occurring in an Alfalfa Field Near Pomona^ California
Method k 95 Confidence Limits
I. FREQUENCY OF CAPTURE
1. Poisson, Both Sexes, 1st Day.
Z. Same as Preceding, Except: Znd Day
3. Same as Preceding, Except; Based
on Females Only, 1st Day; Results
Were Doubled (to Include Males).
4. Frequency of Capture - Geometric
Model, Both Sexes, 1st Day.
5. Same as Preceding, Except: Based
on Females Only; Results Were
Doubled (to Include Males).
6. Frequency of Capture - Regression
Method, Both Sexes, 1st Day.
7. Same as Preceding, Except: Based
on Females Only, 1st Day; Results
Were Doubled (to Include Males).
239
185
293
46
Not
Calculated
274 204 44
401 No Procedure Available
442 No Procedure Available
290 Not Calculated
315 Not Calculated
II. TOTAL COUNTS ON SAMPLE PLOTS
1. 1st Day 87
2. Same as Preceding, Except: 2nd Day. 102
III. RELATION OF VARIANCE TO MEAN
1. Cursory Coiints, 111 Plots Occurring in
8 Rows, 1st Day. 0
2. Same as Preceding, Except: 2nd Day. 0
3. Same as Preceding, Except: Based on
Sum of Each of 8 Rows; 1st Day. 664
4. Same as Preceding, Except: Data
for Both Days Combined (n = 16 Rows). 0
5. Same as Preceding, Except: Whole
Field Subject to Scanning, 3rd Day. 0
IV. MARK-RATIO MODEL
65 109
Not Calculated
Not Calculated
Not Calculated
21
Not Calculated
Not Calculated
1, The Dahl, or Petersen Method, Data 307 210
from Both Days
610
V. REMOVAL METHOD
i. Data from 1st Day.
187
0
402
7 (1): 35-49, 1968
POPULATION ESTIMATORS
43
data for plotting the regression line is obviously subject to all
of the problems affecting other methods based on marked ani-
mals (see Ricker, 1958:86-100; Hanson, 1967).
TOTAL COUNTS ON SAMPLE PLOTS
On the first day 87 animals were counted and on the second
day two counts yielded, respectively, 87 animals (Table 3) and
102 animals. These figures were undoubtedly much too low,
mainly because resting butterflies were tending to fly off the
plots before the observers could determine whether the butter-
flies were within a plot’s boundaries. Also, the relatively rapid
and erratic flight paths of moving butterflies made it difficult
to tell when they were above a given plot and the observers
erred on the conservative side. Since the confidence limits for the
first day’s estimate were rather narrow (Table 3), the bias ap-
peared to be rather consistent. The method of total counts could
be made more useful by (a) marking off the boundaries in a
more elaborate, easily-recognized way than was done here and
(b) by enlarging the plots; but we believe that, for highly
mobile animals such as Colias eimjtheme, the plotless, frequency-
of-capture methods are superior when ingress and egress are not
important problems.
RELATION OF VARIANCE TO MEAN
• This method is described (Hanson and Chapman, in press;
Hanson, 1967) as a method for rapidly estimating the number
of groups of free-ranging animals from cursory, incomplete
counts. None of the animals need be marked or removed, and
total counts of any component are not required; but, on the
other hand, the model requires ( in addition to the usual random
sampling) that the data come from a binomial distribution.
Although individual animals usually tend to be clumped spa-
tially, the groups themselves should be distributed more at ran-
dom, leading to a binomial distribution of groups. Therefore,
the model deals only with groups. After the worker estimates
the total number of groups, he would of course multiply by
the average group size to get total population. Since the alfalfa
butterflies were here essentially solitary, except for some very
brief liaisons between copulating individuals, it turned out that
group size was usually 1. However, as is indicated by the esti-
mates shown in Table 3, the proper data could not be obtained.
The data on the counts of individuals seen per plot and the
resulting variance and mean per plot are shown in Table 2.
When the data were substituted in the proper formula the results
gave
NUMBER OF ANIMALS
44
HANSON-HOVANTTZ
]. Res. Lepid.
NUMBER OF CAPTURES PER INDIVIDUAL
7(1): 35-49, 1968
POPULATION ESTIMATORS
45
.784-^
K == =0
.784 1.116
Since the variance exceeded the mean, this caused a negative
estimate, interpreted biologically as a population of size zero
(Table 3).
When the preceding samples of Table 2 were combined within
each of the 8 transects to smooth out random error, 8 samples
of butterflies were obtained: 8, 17, 11, 7, 11, 12, 13, 8. For this
series, the mean was 10.875 and the variance was 10.697, leading
to the following estimate of the total population (Table 3):
K = 10.8752 / (10.875 — 10.697) = ca. 664.
The 90% confidence limits were obtained from Dr. Chapman’s
equations (Hanson and Chapman, in press)
1 — (7) (10.697) < p < 1 - (7) (10.697)
(10.875) (2.17) (10.875) (14.1)
where 2.17 and 14.1 are the upper and lower values of
Chi-square, for 7 d.f. and .95 and .5 probability, respectively,
read off from a table such as that of Fisher and Yates (1957:45).
After the indicated arithmetic is performed, it resulted in
1 _ 3.17 < p < 1 __ .488 .
Since a negative value of p in this double inequality (on the
left alone) is biologically impossible, the lower bound could not
be less than 0, and the confidence interval for the probability of
seeing a given animal became
0 < p < .512 .
The confidence limits for K finally became 10.875 / .512 = 21.24;
and 10.875 / 0, which can be taken as infinity.
All other attempts to estimate K from the relation of variance
to the mean failed because the variance was too high. Evidently
(a) the true population density varied greatly from one plot to
the other or (b) the animals were aggregated into larger group-
ings that were not recognized as such, or (c) the activities of the
observer introduced considerable extraneous variation. Most
likely each problem occurred to a degree.
First, the outside transect on each side of the field appeared
to continually have fewer butterflies than did the inner transects;
why the butterflies tended to use the outside parts of the field
less, was not clear, but superficially the alfalfa appeared thinner
there.
Second, at times the butterflies were momentarily aggregated
a female, but these groups were treated as chance events and the
46
HANSON-HOVANITZ
J. Res. Lepid.
Colias in them were recorded as individuals (i.e., several
“groups” containing one animal each).
Third, the principal cause of the excessive variation seemed
to be the lack of an objective method for determining the bound-
aries of the area scanned and whether or not observed butter-
flies were within those boundaries during the rapid, cursory
counts. Since the estimator based on relations of the mean to
the variance would provide an easy and rapid way of estimating
density if the proper data can be obtained, it is important to find
an objective way to make the counts.
MARK-RATIO MODEL
The well-known mark-and-recapture method, apparently first
used on animals by Dahl (1917), and reviewed extensively by
Ricker (1958), Southwood (1966), and Chapman (1954), was
tried here; for data we had 128 different butterflies caught the
first day and 24 caught the second day, of which 10 had been
marked at least once. Therefore,
, (128) (24)
K = — 307 ,
10
with limits (210, 610) (Table 3).
The small size of the sample caught on the second day, small
in spite of considerable effort, indicated that much of this Colkis
population had left the field. Egress would cause no problem so
long as the ratio of marked to unmarked animals did not change.
Since there seemed to be no evidence that marked animals were
leaving at a faster r^ite than the others, the estimate of 307 was
reasonably close to the true population size. The confidence
limits were somewhat wide, mainly because of the small sample
in loose groups, perhaps due to attraction of several males to
size collected on the second day.
REMOVAL METHOD
The removal method of population estimation was apparently
begun by Hjort and Ottestad ( 1933 ) and has since been re-
viewed by several persons, including particularly Zippin (1956).
In the present work it was expected that the count for the first
day could be compared to that for the second day although no
animals nor plots were removed. It was planned that any animal
caught on the second day that bore a mark from the first day
would be treated mathematically as dead. However, the decline
in population size during the two days negated one of the main
requirements for use of the removal models. As a result, another
approach was tried.
7 (1): 35-49, 1968
POPULATION ESTIMATORS
47
For both sexes combined, 81 butterflies were caught once on
the first day, and 35 were caught twice on that day (Table 1).
Now let it be imagined that two independent samples had been
taken on that day, each involving equal effort and the other
standard assumptions of the removal method, and that in the
first sample 81 animals were caught. If efforts, etc., were con-
stant, then 81 should have been caught in the second (hypothet-
ical) sample, of which 35 would have been carrying earlier marks.
The 35 marked ones (Table 1) found in the second imaginary
sample may be subtracted from the 81 assumed caught, leaving
46 as the size of the unmarked portion in the second sample.
This manipulation provides the raw data for use in the estimat-
ing equation:
812
K = ,= 187 .
81 — 46
Where Ci and Co are the number of animals caught and removed
on the first and second surveys, respectively, then the standard
error of the estimate is (Zippin, 1956)
cfc| (c^ -f c^) (6561) (2116) (127)
IQS .
(ci ~ C2)' 1,500,625
Therefore, the upper and lower limits, at the 95% confidence
level, were (0, 402) (Table 3).
The estimate of the total population size, 187, seemed too
small, although the confidence limits included the most reason-
able values, 275 to 300. The difficulty seemed to be that more
unmarked animals should have appeared in the second sample,
requiring that the number marked for the second time should
have been smaller. Therefore, the possibility was present that
once a butterfly was marked, it was more prone to be caught
again, but if this were so, the estimate based on the Dahl mark-
ratio method should have been smaller. The question was not
definitely answered but “proness to capture” should not have
caused much trouble.
DISCUSSION AND SUMMARY
How satisfactory any estimator of density does perform de-
pends in part on each person’s concept of what is “satisfactory.”
According to our experience, most zoologists expect results too
close to the real population mean and often seem to think that
an error much over 10-20% is excessive. Yet considering the many
possible sources of error even in stationary populations such as
48
HANSON-HOVANITZ
/. Res. Lepid.
plants, it is a wonder that a highly mobile animal group can
have its density estimated within one order of magnitude. Cer-
tainly it appears that estimates on highly mobile animals should
be considered reasonably good if they are within 50% of the true
population size, although attempts should of course continue to
be made to find better techniques.
Viewed in this light, several estimates obtained in the present
study were fairly close to what seemed reasonable, that is about
275 to 300; frequency-of-capture models, based on either the
Poisson distribution or on a regression line, and the mark-ratio
model gave estimates near that value. Methods based on finding
plot boundaries, such as the mean-variance model or total counts,
were not as satisfactory, although they might become so when
the plots are larger and better marked. At least 128 different
butterflies were caught and marked, and the latter sets a known
minimum limit for the population. The upper limits were either
about 344 (frequency-of-capture, Poisson), 402 (removal meth-
od), or 610 (mark-ratio method) (Table 3). Which of these is
better cannot be dogmatically stated, since the correct answer
rests partly on a matter of intuition, and confidence level associ-
ated with the value selected. In our opinion, the true upper
limit of the population estimate should have been not more
than about 400, i.e, 25% above the upper end of the most prob-
able estimate of K.
The only adequate method for deciding the proper size of K
and its confidence limits is to repeat the experiment a number
of times, within a fairly short interval of time, using a variety
of models, and particularly obtaining the basic data by a variety
of field methods. Unfortunately, if such intensive efforts had been
made here they would have driven even more of the butterflies
from the place of study, and excessive egress was already the
principal difficulty in the present work. Therefore, continued
research should be done to find additional methods for estimat-
ing density, particularly ones that disturb the population a
minimum. The model recently proposed by Hanson (1968)
might be helpful in this regard. In a nutshell, the best suggestion
for lepidopterists, and zoologists in general, seems to be that
they should use several good methods on each population stud-
ied and be prepared to accept errors up to 50% of the estimates .
made.
7(1): 35-49, 1968
POPULATION ESTIMATORS
49
ACKNOWLEDGMENTS
Wish to thank the following persons: Robert T. M'Closkey,
Eric Hovanitz and Roderick Hanson for help with the field;
Professor Paavo Voipio, in whose Institute of Zoology of the
University of Turku, Finland, the data were analyzed and most
of the paper was written; and Miss Linda March who assisted
in typing the manuscript.
LITERATURE CITED
BAILEY, N. T. J, 1959, Statistical methods in biology. John Wiley & Sons,
Inc.: New York, 200pp.
CHAPMAN, D. G. 1954. The estimation of biological populations. Ann.
Math. Statistics, 25:1-15.
CRAIG, C. C. 1953. On the utilization of marked specimens in estimating
populations of Hying insects. Biometrika, 40:170-176.
DAHL, K. 1917. Studier og Forsok over Orret og Orretvand. Kristiana,
Norway, 182pp.
EDWARDS, W. R. and EBERHARDT, L. 1967. Estimating cottontail
abundance from livetrapping data. /. Wildl. Mgmt., 31(1):87"96.
FISHER, R. A, and YATES, F, 1957. Statistical tables for biological,
agricultural and medical research. (5th ed. ) Oliver and Boyd: London,
138pp.
HANSON, W. R. 1967. Estimating the density of an animal population.
/. Res. Lepid., 6( 3 ) :203-247.
HANSON, W. R, 1968. Estimating the number of animals: A rapid method
for unidentified individuals. Science, 162:675-676.
HANSON, W. R. and CHAPMAN, D. G. Date? A model for rapidly esti-
mating abundance among unidentified animals. In press.
HJORT, J., JAHN, G. and OTTESTAD, P. 1933. The optimum catch.
Hvalradets Skrifter, 7:92-127.
HOVANITZ, W. 1948. Differences in the field activity of two female color
phases of Colias butterflies at various times of the day. Contrib. Lab.
Vert. ZooL, Univ, Michigan. 41:1-37.
RICKER, W. E. 1958- Handbook of computations for biological statistics
of fish populations. Queen’s Printer; Ottawa, Canada, 300pp.
SOUTHWOOD, T, R. E. 1966. Ecological methods, with particular refer-
ence to the study of insect populations. Methuen & Co., Ltd.:London,
391 pp.
STEEL, R. G. D. and TORRIE, J. H. 1960. Principles and procedures of
statistics with special reference to the biological sciences. McGraw-
Hill: New York, 481pp
ZIPPIN, C. 1956. An evaluation of the removal method of estimating animal
populations. Biometrics, 12(2): 163-189.
50
HOVANITZ
/. Res. Lepid.
HABITAT: ARGYNNIS CALLIPPE LAURINA
The iinsilvered form of this complex which includes calUppe,
comstocki, raacaria, lamina^ coronis, rupestris and others is found
along the lower elevations of the western side of the Sierra
Nevada mountain range in California. Occasional specimens of
wholly or partially silvered are to be found throughout the range
of the race but they are most common toward the south where
populations gradually increase in the percentage of silvering
until the populations may be called rnacaria in the Kern basin.
Silvering picks up again at the north end of the range.
At this location, the primary local vegetation is the Digger
Pine and oak woods. Adults fly after the grass is dry, late June
and early July. The photograph was taken a few miles southeast
of Mariposa, Mariposa Co., California, early July, 1969.
William Hovanitz
Journal of Research on the Lepidoptera
7 (1): 51-55, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A.
© Copyright 1968
ON MEXICAN SATYRIDAE,
WITH DESCRIPTION OF A NEW SPECIES
LEE D. MILLER
The Allyn Foundation,
Rm. 885, 222 West Adams Bldg.,
Chicago, Illinois 60606
The Carnegie Museum-Catholic University of America
expedition to eastern Mexico in January, 1966, collected relatively
few Satyridae, mostly of rather common species. Two specimens
were of special note, and these are recorded here.
A single female of Dioriste tauropolis (Westwood) was col-
lected on 9 January 0-3 miles northwest of Gomez Farias, Ta-
maulipas, between 280 and 700 m. elevation in the tropical ever-
green forest. This specimen and another taken by Mr. L. I. Gil-
bert (personal communication) apparently represent the furthest
north records for tauropolis. Godman and Salvin (1879-1901:
108) report it from Cordova and Jalapa, Veracruz, while Hoff-
mann (1940: 670) lists this species from only Veracruz, Tabasco,
Chiapas and Oaxaca. The present specimen (L. D. Miller speci-
men no. 1966-373) was colelcted flying in the sunlight along a
woodland road.
Members of the genus Cyllopsis infrequently are collected in
series, so it was a pleasant surprise to take a series of nine
specimens in a semi-montane situation east of Ciudad Victoria,
Tamaulipas. When the material was prepared eight of the
specimens were of the realtively common C. gemma freemani
(Stalings and Turner), but the ninth was totally unlike any
Cyllopsis I have seen. My first impression was that the specimen
was a singular aberration, but its genitalia are totally unlike
those of freemani, and this specimen apparently represents a new
species.
51
52
MILLER
/. Res. Lepid.
Figs. 1-2. Cyllopsis dospassosi, new species, Holotype $ . Fig. 1, upper
surface. Fig. 2, under surface. About 3.5 times natural size.
7(1): 51-55, 1968
MEXICAN SATYRIDAE
53
Cyllopsis dospassosi, new species
Figs. 1, 2 (Holotype ^ ), 3 (genitalia of Holotype $ )
Male: — Head, thorax and abdomen dark brown densely
clothed with dull brown hairs above and tan ones below. Palpi
dark brown densely clothed with dark brown hairs above and tan
ones below. Antennae dark brown dorsad, tan ventrad with the
shaft ringed in dark brown. The legs are clothed with dense
tan hairs. Forewings above dull grayish-brown shading to
darker dull brown marginally and around the apex to about
halfway down the costa; otherwise unmarked. There is no
androconial patch of mealy scales below the cell, as are shown
in the hilaria (Godman) group. Hindwings above also dull
grayish-brown, darker at the apex, with double blackish-brown
marginal spots in spaces M2-M3 and M.o-Cui, as well as a faint
smaller single spot in Cui-Cus, and the bands of the under side
showing through faintly. Forewings beneath olive-tan marked
with olive-brown as follows: many scrawls in the basal part of
the cell, a large patch at the end of the cell, a spotband just
outside the cell from the apex to 2A, the spot in 2A being doubled
with the proximal member situated directly below the cell spot,
and a marginal spotband from cells Rr,-Mi to Cu2-2A. The hind-
wings below are of the same olive-tan color as the forewings,
scrawled basally with olive-brown and with two transverse
broken bands of the same color from the costa to near the inner
margin, one across the cell and the other outside it, and with
two silver-centered dark brown ocelli marginally in spaces M2-
M.i and M.rCui along a thin, wavy, silver marginal band ex-
tending from spaces Rs-Mi to Cu-Cu2. The fringes are uniform-
ly dull brown above and tan beneath on both wings. The length
of the forewing of the Holotype $ is 16 mm.
The male genitalia bear little resemblance to those of freemani
but are rather close to those of hilaria, especially as regards the
straighter uncus and the blunter valvae. The valvae of freemani
are tapered to a point (Fig. 4). For comparison the valva of
hilaria is shown in Fig. 5.
Female: — Unknown.
Holotype $ : — 52 mi. E. of Ciudad Victoria, Tamaulipas,
MEICO, 510 m., 7 Jan. 1966 (C. M. — C. U. A. expedition); L.
D. Miller specimen no. 1966-119; S genitalia slide no. 1496 (Lee
D. Miller). The Holotype is in the collection of Carnegie Mu-
seum.
7 (1): 51-55, 1968
MEXICAN SATYRIDAE
55
I take great pleasure in naming this distinctive species for
Dr. C. F. dosPassos of Mendham, New Jersey, who was in part
responsible for the expedition. His work has put him in the
forefront of American lepidopterists.
Cyllopsis dospassosi is totally unlike freemani, pyracmon
( Butler ) , or any of the other species previously recorded from so
near the United States, as shown by both the pattern and the
genitalic structures. The present species is most closely allied
to hilaria, pephredo (Godman) and their relatives, but it may be
distinguished immediately by the grayer upper surface and the
olive-tan under surface with no trace of the rust color that char-
acterizes the rest of the hilar ia- group. The male genitalia differ
in only minor respects from those of hilaria^ but such close cor-
respondence in the terminalia of closely related species is the
rule in Cyllopsis. The Holotype of dospassosi was collected in
the tropical deciduous forest perched in a brush pile at the edge
of the dense woods. This species should be sought in other
suitable habitats in Tamaulipas, particularly further south in
the Sierra de Tamaulipas proper, and in the low coastal hills
of northern Vercruz. There is a remote possibility that dospassosi
may be found in southern Texas if suitable habitat for it can
be found.
LITERATURE CITED
GODMAN, F. D., and O. SALVIN, 1879-1901. Biologia Centrali-Ameri-
cana. Insecta: Lepidoptera-Rhopalocera. London, 2 vols.
HOFEMANN, C. C., 1940.
lepidopteros mexicanos.
Biol. 11: 639-739.
Catologo .sistematico y zoogeografico de los
Primera parte. Papilionoidea. An. Inst, de
56
HOVANITZ
/. Res. Lepid.
HABITAT: PIERIS BECKERI
Pieris heckeri has more than one food plant. In certain areas in the
Great Basin and in southern California the larvae are restricted to mustards
but in other areas, they are restricted to a member of the Capparidaceae,
namely, Isomeris arhorea. The ability of this plant to remain green when
all else has dried up probably accounts for the existence of adults of this
species flying in the dry washes and hills of desert habitats late in the
season.
The photographs were taken near Caliente, Kern Co., Calif, early
July, 1969.
Journal of Research on the Lepkioptera
7 (1): 57-63, 1968
1140 W. Orange Grom Ave., Arcadia, California, U.S.A.
© Copyright 1968
IDENTITY OF THE MOTH
“STRETCH I A" BEHRENSIANA (GROTE)
WITH NEW SYNONYMY
(NOCTUIDAE)
JOHN S. BUCKETT
Systematic Entomologist,
Bureau of Entomology,
California Department of Agriculture,
Sacramento, California
Until present times, “Stretchid' Behremiana (Grote) has
been considered a mystery. It was the author’s intention to bring
this matter into print years ago, but the manuscript was mis-
placed until recently.
“Stretchia” behremiana is a name that is representative of an
uncommon form of the species concerned, and as Orthosia
macona (Smith) is the more recent name representing the same
entity, it will have to fall into synonymy of behremiana. The
primary clue which led to the proper identification of behrem-
iana is the colored illustration presented by Hampson (1905).
Until this colored illustration of the type female (pL 89, fig. 10)
of behremiana was properly associated, the name had been ap-
plied to several genera, none of which are correct by present day
concepts. The moth concerned is a typical Orthosia^ and there-
fore it is placed as a new combination within Orthosia.
In the past few years, a number of specimens have been col-
lected in central California that match Hampson s (op. cit. )
colored illustration of the type. Illustrated by figures 1-4, one
can see the major range in the maculation of the primaries of
behremiana. Under the redescription, the variation in coloration
of the imago is given.
57
58
BUCKETT
J. Res. Lepid.
Fig. 1 Orthosia behrensiana ( Grote ) , female. Anderson Springs, 4 miles
northwest of Middletown, Lake County, California, 21 February 1954
(W. R. Bauer). This specimen matches type and is in the collection
of the British Museum of Natural History, London.
Fig. 2 O. behrensiana, female. Cobb Mountain, Lake County, California,
12 February 1955. (W. R. Bauer and J. S. Biickett). This specimen
also in British Museum of Natural History.
7(1): 57-63, 1968
\STRETCHIA” BEHRENSIANA
59
Orthosm hehrensiana (Grote), New Combination
Graphiphora Behremiana Grote, 1875. Canad. EntomoL 7(4) :71-
72.
Perigrapha behrensuma^ Grote, 1881. Canad. Entomol, 13(6) :133;
Smith, 1889. Proc. United States Nat. Mus. 12:493-494,
Stretchia behremiana, Smith, 1891. Trans. American Entomol
Soc. 13:120- 1893. Bull United States Nat .Mus. No. 44,
p.208; Dyar, 1903 (1902). Bull United States Nat. Mus.,
No. 52, p.l67; Woodworth, 1912. California Monthly Bull
1(10): 789 (indicates Xylomania, after Hampson's concept);
McDunnoiigh, 1938. Mem. Southern California Acad. Sci.
1:75.
Xylomania behremiana, Hampson, 1905. Cat. Phalanae British
Mus. 5:390 plus pi 89, fig. 10; Woodworth, 1912. California
Monthly Bull 1(10): 789 (cites as a Stretchia, in italics).
Xylomiges behremiana, Barnes and McDunnough, 1917. Check
list of the Lepidoptera of Boreal America, p.53; Draudt (in
A. Seitz), 1923. The Macrolepidoptera of the World, Div.
2, vol 7, p.l5, pi 22, row f.
Taeniocampa macona Smith, 1908. Ann. New York Acad. Sci.
18(2), part 2: 102-103; Rindge, 1955. Bull American Mus.
Nat. Hist. 106(2): 119.
Orthosia macona, Barnes and McDunnough, 1917. Check list of
the Lepidoptera of Boreal America, p.54; Draudt (in A.
Seitz), 1923. The Macrolepidoptera of the World, Div. 2,
vol 7, p.l59; McDunnough, 1938. Mem. Southern California
Acad. Sci. 1:76. New Synonymy.
Male: Ground color of primaries dorsally varying from pale
tan to dark brown; secondaries dorsally off-white, iorrated with
varying degrees of dark brown scales.
Head with vertex clothed in admixture of tan and dark brown
elongate simple hairs; frons clothed in almost uniformly tan col-
ored elongate simple hairs, integument evenly truncately round-
ed; labial palpi exterolaterally clothed in blackish scales, and
hairs, ventrally clothed in elongate tan and sparsity of black
simple hairs terminal segment minute; antennae with scape and
pedicle clothed in short, broad white to tan scales; flagellomeres
biserrate-fasciculate, serrations becoming less pronounced term-
inally, dorsally clothed in tan simple, dentate scales.
Thorax with collar nearly unicolorous tan, composed of elongate
simple hairs, terminal segment minute; antennae with scape and
60
BUCKETT
/. Res. Lepid.
Fig. 3 O. hehrensiana, male. Siimmerland, Santa Barbara County, Cali-
fornia, 24 January 1948 (C. W. Kirkwood).
Fig. 4 O. hehrensiana^ female. Ojai, Ventura County, California, 16 June
1957 (W. E. Simonds).
7 (1): 57-63, 1968
“STRETCHIA^’ BEHRENSIANA
61
clothed anteriorly in blackish and tan elongate simple hairs,
posteriorly clothed in oflF- white to tan elongate simple hairs;
prolegs with femur and tibia clothed intero-laterally in smoky
and tan simple scales and hairs; tarsus clothed in dark scales,
terminally each tarsomere clothed in tan scales; meso- and meta-
thoracic femora and tibiae clothed in tan elongate hairs and
scales, extero-laterally with admixture of dark brown simple
scales, tibial spurs one and two are meso- and metatibiae, respec-
tively; primaries dorsally with pale tan to dark brown ground
color; basal half line occasionally represented in either black
or tan; transverse anterior line when present, geminate, central-
ly lighter than ground color, direction as in fig. 1; orbicular,
when obvious, subcircular, may coalesce with reniform, tan or
orangish-tan; median area of ground color, or median shade may
be present (as in fig. 2); reniform suboval, smaller end pointed
toward costa or apex, colored as in orbicular; transverse posterior
line concolorous with transverse anterior line, direction as in fig.
1; subterminal space of ground color; subterminal line irregular
in course when discernable, lighter than ground color; terminal
line represented by black dots between veins; fringes more
yellowish than ground color; ventral surface brownish, iorrated
with black; reniform represented by black splotch; transverse
posterior line brown; veins between transverse posterior line
and fringes slightly outlined in tan; terminal line as in dorsal
surface; secondaries dorsally off-white, basally slightly darker
than apically; distal dot dark brown; terminal line dark brown
(as in figures 1, 2, and 4); ventrally as in dorsal surface, except
costal area darker.
Abdomen dorsally clothed in admixture of dark brown and
tan simple scales; terminally clothed in tan elongate scales and
simple hairs. Genitalia as in figures 5 and 6.
Greatest expanse of forewing 14-17mm.
Female: As in male, except for antennae which are ciliate,
fasciculate; tendency for the '‘behrensiana form” by far greatest
in this sex. Greatest expanse of forewing averaging slightly
larger.
O. hehrensiana is quite a variable species in dorsal maculation
of the primaries. For this reason the correct name of the entity
in concern has remained in confusion until recently. The species
is widespread in California, ranging from San Diego County
north into Del Norte County, and from sea level into the Sierra
Nevada Mountains.
62
BUCKETT
/. Res. Lepid.
Fig. 5 O. hehrensiana, male genitalia minus aedeagus; Bauer-Biickett
Slide No. 68B27-50. Anderson Springs, Lake County, California, 21
March 1949 (W. R. Bauer).
Fig. 6 O. hehrensiana, aedeagus of male genitalia. Data same as fig. 5.
7 (1): 57-63, 1968
“STRETCHIA” BEHRENSIANA
63
Specimens of “macond" were sent to Dr. F. H. Rindge of the
American Museum of Natural History, New York, where he
graciously made type comparison with Smith’s type of T. ma~
cona. The conspecificity of the specimens sent by the author
with the type was confirmed.
Specimens of behremmna were also sent for type comparison
to Dr. I. W, B. Nye of the British Museum of Natural History,
London. Unfortunately, the type female is lacking an abdomen,
so genitalic comparison was impossible at this time; the type is
otherwise in excellent condition.
I would like to extend my appreciation to both Dr. Rindge and
to Dr. Nye for their cooperation in this project, and to Mr.
■George M. Buxton of this Buerau for the photographs contained
herein.
LITERATURE CITED
BARNES, WM., and J. H. McDUNNOUGH, 1917. Checklist of the Lepi-
doptera of Boreal America. Herald Press, Decatur, Illinois, 392 +
viii pp.
DRAUDT, M. (in: A. A. Seitz), 1923. The Macrolepidoptera of the
World Alfred Kerean Press, Stuttgart. voL 7, 412pp. + 96 pis.
DYAR, H. G., 1903. (1902). A list of North American Lepidoptera and
key to the literature of this order of insects. Bull. United States Nat.
Mus., No. 52, xix -f 723pp.
GROTE, A. R., 1875, Preliminary list of the Noctiiidae of California.
Canad. EntomoL 7(4):67-72.
^ 1881 New Noctuidae ,with a list of the species of Perigrapha.
Canad. EntomoL 13( 6 ): 131-134.
HAMPSON, G. F., 1905. Catalogue of the Noctuidae in the collection of
the British Museum. Longmans & Co., London, England, vol. 5, xvi
-|- 634pp.
McDUNNOUGH, J. H., 1938. Check list of the Lepidoptera of Canada
and the United States of Aemica, part I, Macrolepidoptera. Mem.
Southern California Acad. Sci., vol. 1, 272pp.
RINDGE, F. H., 1955. The type material in the J. B. Smith and G. D,
Hulst collections of Lepidoptera in. the American Museum of Natural
History Bull. Mus. Nat. Hist., 106( 2 ) :91-172.
SMITH, J. B., 1889. Contributions toward a monograph of the Noctuidae
of temperate America — Revision of some taeniocampid genera. Proc.
United States Nat. Mus. 12:455-496 + pks. 22 and 23.
, 1891. Notes on some Noctuidae, with descriptions of new genera
and species. Trans. American EntomoL Soc. 13:103-135 + pi. 2.
1893. Catalogue of the lepidopterous siiperfamily Noctuidae
found in Boreal America. Bull. United States Nat. Mus. No. 44, 424pp.
^ 1908. New species and genera of the lepidopterous family Noc-
tiiidae for 1907, part 2. Ann. New York Acad. Sci. 18(2) :91-127.
WOODWORTH, C. W., 1912. Check list of California Insects II. California
Monthly Btdl 1( 10) :782-790.
NOTICES
BOOKS;
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastimil Choc. Paul Hainlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophical
Library, N.Y.
WANTED:
Brephidium exilis, B. fea, B. isophthalma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, California 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systematics^ biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
Manuscripts for immediate publication in this JOURNAL. With color
nlay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3x5 card in the form desired and they will be printed in the
next following issue of the JOURNAL.
THE J0UR.HAL
Volume 7 Number 1 March, 1968
IN THIS ISSUE
Controlled environment experiments with
Precis octavia Cram. L. McLeod 1
Ecological and Distributional notes on Erehia disa
in central Canada John H. Masters 9
Methods for Studying the Chromosomes of Lepidoptera
Thomas C. Emmel 23
Note on Vital Staining of Actias luna silk
John M. Kolyer 29
Present and Ice Age Life Zones and Distributions
W. Hovanitz 31
Trials of several density estimators on a butterfly
population W. R. Hanson and W. Hovanitz 35
Habitat — Argynnis caUippe latirina
W. Hovanitz 50
On Mexican Satyridae Lee D. Miller 51
Habitat — Pieris beckeri ‘W. Hovanitz 56
Identity of the moth “Stretchm’ behrensiana
with new synonymy. J. S. Buckett 57
5 ^5 * 7^^
tyifl
vz
t
published by
The Lepidoptera Research Foundation, Inc.
at
1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz
Associate Editors:
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Florida 32601.
Maria Etcheverry, Centro de Estiidios Entomologicos, Casilla 147, Santiago,
Chile.
T. N. Freeman, Div. of Entomology, Dept, of Agriculture, Ottawa, Ontario,
Canada.
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G. de Lattin, Zoologisches Institut, Universitat des Saarlandes, Germany.
Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.
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Illinois 60181.
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THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a
year, Spiing (March), Summer (June), Autumn (September), and Winter (December)
by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
cation and the general business office are located at 1160 W. Orange Grove Ave.,
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Journal of Research on the Lepidoptera
7(2) : 65-86, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
STUDIES ON NEARCTIC Euchloe,
PART 5. DISTRIBUTION.
PAUL A. OPLER
University of California,
Berkeley, California
Populations of the Nearctic species of Euchloe occur from
north of the Arctic Circle in Northwest Territories south to Baja
California del Norte, Arizona, and New Merico (30° North
Latitude). In the west they occur to about 150° West Longitude
in Alaska and to 80° West Longitude in Maryland to the east.
The four species of Nearctic Euchloe are single-brooded with
one possible exception. Adults fly as early as January in San
Diego County, California and as late as mid-August at high
altitudes in Colorado.
Within the wide geographic and seasonal parameters men-
tioned above each species has narrower limits. The populations
of each species possess flight period, behavior pattern, and host
plant specificity characteristics that are met by only a small
proportion of possible habitat spaces and times of the year. In
this paper, I will summarize the information bearing on the
distribution in time and space of Euchloe creusa, Euehloe hyan-
tis, and Euchloe ausonides. A later paper will give in detail the
distributional features displayed by populations of Euchloe
oltjmpia (Clench and Opler).
SEASONAL DISTRIBUTION
Populations of Euchloe ausonides found in the lowlands of
central California appear to be at least partially double-brooded
as has been shown by Sette (1958). The basis for this assump-
tion of bivoltinism is the bimodal distribution for a large number
of adults plotted against date of collection. Collection records of
317 individuals collected in the counties bordering San Fran-
cisco Bay are summarized in Figure 1. The small “second
brood” can be produced due to the permissive climate and pres-
65
r^ARCH APRIL MAY I JUNE JULY
66
OPLER
J. Res. Lepid.
1
I
c
Fig. 1 Seasonal distribution of Euchloe ausonides in the San Francisco
Bay Area, California. Solid portion of each bar represents number of males;
open portion of each bar represents number of females.
7(2) : 65-86, 1968
NEARCTIC EUCHLOE
67
ence of suitable host plants over a long period of time. Pupal
diapause, which is normally obligatory for Nearctic Euchloe,
must be facultative to allow those “second brood” individuals
which do occur to complete development in the same year.
All other populations of Euchloe in the Nearctic Region are
clearly single-brooded. The temporal distributions of all popu-
lations are determined by physical permissiveness of the en-
vironment and are coincident with the presence of appropriate
cruciferous host plants in a suitable state of growth to serve as
oviposition sites.
It is interesting to note the displacement of flight periods that
occurs where Euchloe ausonides occurs in sympatry with or in
proximity to each of the other three species. Populations of both
E. creusa and E. hyantis can be seen to have earlier flight periods
relative to E. ausonides in most instances. All examples of sym-
patry known to the writer are summarized below. Synchronic
collections are also indicated.
SYMPATRIC OCCURRENCES WITH
EUCHLOE AUSONIDES
Locality
Dates of capture ( inclusive )
Euchloe creusa Other species Synchrony
E. ausonides
Whitehorse, Yukon
Terr., CANADA
vi"9 vi-9-66
v-29 to vii-5
AlCan Highway, MP 855
Brit. CoL, CANADA
vii-1 vii-1-66
vii-1
Atlin, Brit. CoL,
CANADA
vi-7 to vi-28
vi-20 to vii-7
Nordegg, Alberta,
CANADA
vi-5
vi-25
Banff vicinity,
Alberta, CANADA
v-29 to vi-3()
vii-4 to vii-9
Prince Albert,
Sask., CANADA
v-27 to vi-2 v-27-51
vi-2-51
v-27 to vi-2
Euchloe olympia
Clear Ck. Cyn., nr. Golden,
Jefferson Co., Colo,
v-10
v-26
Lefthand Cyn.,
Boulder Co., Colo.
v-30
v-25
Sunshine Cyn.,
v-4
iv-27 to v-5
Boulder Co,, Colo.
68 OPLER /. Res. Lepid.
Euchloe hyantis
Lilooet, Brit. Col,,
Cranbrook, Brit., Col.,
CANADA
iv-2() to iv-28
iv-18 to v-30
vi-18 to vi-25
v-9 to vii-9
Alta Lake,
Okanogan Co., Wash.
iv-29
iv-29-51
iv-29
Baker, Baker Co., Ore.
v-20 to v-21
v-27
Ontario, Malheur Co.,
Ore.
iv-30
iv-30-41
iv-30
Eureka, Juab Co., Utah
v-7 to v-29
v-23
Salt Lake City,
Salt Lake Co., Utah
V-?
iv-18 to vi-13
Jackson Hole,
Teton Co., Wyo.
v-16 to v-23
v-20-24
v-23-24
v-20 to vi-17
Mt. Wheeler,
White Pine Co., Nev.
V- 19/24
v-19/24-29
v-19 to vi-8
2 mi, N Mt. Shasta City,
Siskiyou Co., Calif.
vi-25 to vii-3
vii-3-55
vii-3
Cloverdale,
Sonoma Co., Calif.
iv-25
iv-15
Hallelujah Jet.,
Lassen Co,, Calif.
vi-17
vi- 17-67
vi-17
Mono Lake,
Mono Co., Calif,
vi-17
vi-17-19
vi-17
6 mi. W. Lone Pine,
Inyo Co., Calif.
iv-9
iv-9-60
iv-9
GEOGRAPHIC DISTRIBUTION
The geographic distributions of Euchloe aiisonides, creusa, and
hyantis are presented as figures 2, 3, and 4 respectively. More
specific locality information is given in the citation of collection
data.
Euchloe ausonides, the species with the widest range, occurs
sympatrically with each of the other three species at some points
in their distributions, the instances known to this writer are
listed above. In each case of sympatry some altitudinal or eco-
logical information not shown on collection labels probably
occurs to cause effective allopatry in the sense of the niche
occupied by each species.
EGOLOGIGAL DISTRIBUTION
Although no intensive ecological studies have been conducted
on species of Euchloe several statements on the ecological attri-
butes which appear to be important in determining the presence
or absence of populations of Euchloe can be made.
One apparent common denominator of all populations is that
they occur in open areas. The other important parameter is the
occurrence of populations only where some feature of relief in
the landscape occurs. These two features seem to be required
7(2) ; b5-H6, 19(iH
NEARCTIC EUCHLOE
69
70
OPLER
/. Res. Lepid.
Fig. 3. Distribution of Euchloe creusa. Solid circles represent local-
ities for the nomenotypic race; the half-open circle represents the occur-
rence of a distinctive segregate at Prince Albert, Saskatchewan.
7(2) : 65-86, 1968
NEARCTTC EUCHLOE
71
Fig. 4, Geographic distribution of Eiichloe hijantis. Solid squares
represent localities for the nomenotypic race; half-open squares represent
the Sierra Nevadan populations; half-open circles represent localities for
E. hyantis andrewsi; solid circles represent localities for all populations
tentatively referable to E. hyantis lotta. The two groups of circles circled
in black represent^ the “Mt. Pinos Block segregate” and the “Peninsular
Ranges Segregate.” Collection records, for this species at Cranbrook and
Oliver, British Columbia do not appear on this map.
72
OPLER
J. Res. Lepid.
to serve as keys to behavior patterns displayed by adults and to
prevent loss of contact that might occur in ''unbroken land-
scape.” Thus, some populations appear keyed to "hill-tops,”
others to sunny, open areas on hillsides, and still others to river
courses or dry stream beds. I have observed populations of
Euchloe ausonides or E. hyantis keyed to each of these situations.
The preferred habitat of Euchloe creusa in the vicinity of Banff,
Alberta is on the talus slopes of lateral moraines.
It should be further noted that there appears to be ecological
displacement whenever two species of Euchloe occur in close
spatial proximity. This displacement may take the form of flight
behavior, altitudinal occurrence, or host plant selection differen-
ces or any combination of the three. In areas where species of
Anthocharis and Pieris occur with Euchloe^ very complex inter-
actions of an ecological nature must occur indeed. I believe that
a proper description of the ecological characteristics of the
crucifer feeding species of Pieridae in the Nearctic Region must
await more careful field studies than have been made by this
author.
SPECIMENS EXAMINED
The data for specimens seen during this study to date are
listed below. For Euchloe ausonides in the vicinity of San
Francisco Bay, California and for Euchloe ausonides coloradensis
in Colorado where records are extremely numerous only county
names and ranges of collection dates are given. Collections are
abbreviated as follows: American Museum of Natural History
(AMNH), David L. Bauer, (DLB), California Academy of
Science (CAS), California Insect Survey, University of Califor-
nia (CIS), Glenn Gorelick (GG), John Lane (JL), Nevada State
Museum (NSM), J. W. Tilden (JWT), Los Angeles County
Museum (LACM), Noel La Due (NLD), Oakley Shields
(OAS), Fred Thorne (FT), and Paul Opler (PAO).
LITERATURE CITED
CLENCH, H. K. and P. A. OPLER, in preparation. Bionomics of Euchloe
olijmpia.
7(2):65-H6, 196S
NEARCTIC EUCHLOE
73
Euchloe ausonides ausonides
CANADA. - ALBERTA, vie. Banff, ?, vii-4/9-30 (J. F. May,
AMNH); Calgary, Tdcf, 3$5, vi-1-03, vi-4- ?, vi-6-03, vi--13- ?,
vi-14- ?, vi-20-?, vi-27"?, vii-3-04, vii- 8- 04 (C. Garrett, AMNH);
Carbon, 4cf cf , 3$$, v-31-?, vi-6-?, vi-23-04, vi-ZY-? (C, Garrett,
AMNH); Didsbury, 3d'd', 5, vi-7-?, vi-10-06, vi-16- ? (C. Garrett,
AMNH); Lacombe, Zd cf (J. A. Comstock, LACM); Lethbridge, Id,
vi-3-29 (J. H. Pepper, AMNH); Nordegg, Id, vi-25-21 (J.
McDunnough, AMNH); Waterton Lakes, 25$, vi-27-Z3, vi-21-23.
BRITISH COLUMBIA. Alaska Highway, MP836, 1$, vii-1-66
(J. Lane, JL); Alaska Highway, MP 855, Id, I9, vii-1-66 (J. Lane,
JL)); A tlin. Id, vi-22-? (AMNH); Atlin, Wilson Crk., Id, 1$, vi-20-14,
vi-22-14 (AMNH); Atlin, 3dd, vi- 27- 29, vii-3-29 (G. Swarth, CAS);
Cranbrook, 9dd, v-9-12, v-15-10, v-22-10, vi-12-?, vi-13-17, vi-15- ?,
vi- 27-?, vii-2- ?, vii-9-? (C. Garrett, AMNH); Creston, 2dd,
vii- 4-?, vii-12- ? (AMNH); Lilooet, 1$, vi-18/25-30 (Iferr, AMNH);
Nelson, 2dd, 1$, v-9?, vi-7-? (AMNH); Princeton, 1$, vi-?-43
(T. Menzies, GAS); 25mi. N. Princeton, 3400*, Id, vii-27-66
(J. Lane, JL); Robson, 1350*, 2dd, v-?-34 (AMNH); Robson,
3dd, iv-20-41, iv-30-39, v-31-39 (H. F. Foxlee, AMNH);
Summerland, Id, v-31-? (AMNH^ MANITOBA. Brokenhead, 7dd,
59$, v-22/vi-4-36, vi-3-30 (AMNH, LAGM); Red Rock Lake,
Whiteshell Provincial Park, Id, 3$$, v-24-55, v- 29- 53, vi-13- 54
(C. Bird, AMNH); Sandilands Provincial Forest, 8mi. SE Richer,
6dd, v-28-67 (J. H. Masters, PAO). ONTARIO. 3 mi. E.
Beardmore, 3dd, vi-1-58 (P. D. Syme, AMNH); Ft. Williams,
1$, vi-8-61 (G. Perrons, AMNH); Neebing, Id, ?-?-6l (W. Hartly,
AMNH). SASKATCHEWAN. Nesbit Prov. Forest, Prince Albert,
2dd, 3?$, v-27-51, vi-2-51 (Aschim, AMNH); Rivercourse, 1$,
299, v-24-41, v-25-41 (AMNH). YUKON TERRITORY. Alaska
Highway, MP 1032, 3000', I9, vii-15~66 (J. Lane, JL) ; Alaska
Highway, MP 1105, 2400*, 299 (P. R. Ehrlich, AMNH); Whitehorse,
Id, vii-5-30 (D. Fraser, AMNH), 14dd, I699, vi- 6/9- 23, vi- 27- 33
(J. A. Kusche, AMNH), lldd, 499, v- 29/vii-l- 66 (J. A. Ebner,
PAO).
UNITED STATES. - ALASKA. Alfred Credc Camp, Id,
vii-15-22 (Pope, LACM), Id, I9, vi-4-02 (AMNH); Fort Yukon,
Id, vi-12-17 (CAS), 2dd, 499, vi-?-19 (J. A. Kusche, LACM);
Forty Mile Highway, 44 miles from east end, I9, vii-14-66
(J. Land, JL); Haine's Highway, MP 93, Id, vii- 5- 52 (Car son,
AMNH); McKinley National Park, 27dd, I899, vi-13/21-31
(F. Morand, AMNH LACM); Taylor Highway, Dawson Jet. ,
3500', Id, vii-4-55 (PR Ehrlich, AMNH). ARIZONA. PIMA CO. :
Bear Canyon, Id, iv-9/l7-31 (L. Martin, AMNH). CALIFORNIA.
ALAMEDA CO. : ( many records, see fig. 1). ALPINE CO. r
rocks over Carson Pass, 8400', 2dd, vii-2-62, vii-4-63
(N. La Due, NLD), CONTRA COSTA CO. : ( Many records
74
OPLER
/. Res. Lepid.
see fig. 1), EL DORADO CO. ; China Flat, 5000', Id*, vi-Z8-48,
(W. E. Kelson, CIS), FRESNO CO, : nr. Rae Lakes, 11, 000', Id”,
vii-2Z-35 (C. W. Anderson, CIS). INYO CO.: Alabama Hills,
6 miles west Lone Pine, Icf, iv-9-60 (R, L. Langston, CIS).
LAKE CO.: Clear Lake, 1?, v-17-65 (G. Gorelick, GG). MARIN
CO. : (Many records, see fig. 1). LASSEN CO. ; 1 mi. W.
Hallelujah Jet. , Id*, vi-18-67 (G. Gorelick, GG). MODOC CO.:
Cedar Pass, 6350', vii-15-65 (R. L. Langston, CIS), 1$, vi-Z6-58
(J. W. Tilden, JWT); Willow Ranch, Id-, vii-10/Z0-Z8 (AMNH).
MONO CO.: nr. Monitor Pdss, Icf, vi-Z3-6Z (J. Powell, QS);
Mono Lake, Icf, vi-17-19 (AMNH); Silver Lake, Zd'd', v-17-36,
vi-9-35 (LACM). MONTEREY CO. : 5 mi. S. Big Sur, Zefd, v-10-58
(P. A. Opler, .PAO); Carmel Vy. , Hasting's Reservation, Z55,
v- 10-59 (P, A. Opler, PAO): Carmel Vy. , Tularcitos Ranch, 1$,
iv-Z7-55 (J. Powell, CIS); Chew's Ridge, nr. Carmel, Zd'd*.
vi- 3-31 (LACM). NAPA CO,: Calistoga, 1$, v-6-16 (W. N. W.mle,
CAS); Mt. VeedarRd. , Icf , iv- Z7- 58 (P. A. Opler, PAO); 3 mi.
W. Oakville, 1$, v-30-60 (P. A. Opler, PAO). NEVADA CO. :
Wolf Creek, 1400', 1?, iv-Z3-63 (N. La Due, NLD). PLACER
CO. : 3 mi. W. Auburn, North Fork American River, Zd'd*,
iv-9-57, iv-ZZ-63 (N. La Due, NLD); Lake Tahoe, 1$, vii-Z-30'
(CIS); between Roseville and Rocklin, ZOO', 1$, iv-11-61 (N.
La Due, NLD). PLUMAS CO. : Beckworth Pass, Ic/, vi-18-67
(G. Gorelick, GG); Half Moon Lake, Id", vii-Z-67 (G. Gorelick,
GG), 1$, vi-6-61 IN. La Due, NLD); Coloma Road, south of
Fair Oaks, lOd'd', iii-13-61 (N. La Due, NLD); Fair Oaks, Zd'd”,
iii-Z4-59, iv-16-58 (N. La Due, NLD); south of Fair Oaks, 5d'd',
3$$, iii-lZ-6Z, iii-14-6l, iii-ZO-61, iii-Zl-60, iv-9-6Z, iv-15-58
(N. La Due, NLD); 7 mi. S. Isleton, Id", iv-1-50 (R. L. Langston,
RLL); ; North Sacramento, Z55, iv-5-60, V-Z6-59 (N. La Due,
NLD); Rancho Cordoba, 5cf cf^ iv-60 (L. C. Clarke, PAO). SAN
FRANCISCO CO.: ( many records, see fig. 1). SAN JUAQUIN
CO.: 5 mi. SE Tracy, 1$, v-36-44 (R. Smith, CIS). SAN MATEO
CO. : ( many records, see Fig. 1). SANTA CRUZ CO. : Santa ;
Cruz, Id, 455, iii-3-31, iv-7-38, iv-10-38, vi-8-31 (W. C.
Wood, AMNH). SISKIYOU CO. : Gazelle, Z$$, vi-Z4-58 (F.
Powell, CIS); Shasta Mdws. , Z mi. N. Shasta City, 3d'd', vii-
3-58 (N. La Due, NLD). SOLANO CO.: Glen Cove, Carquinez
Strait, 3d'd', vi-Z-66, vi-5-65 (R. L. Langston, PAO, CIS);
Vallejo, Zd'd", 3$$, vi-Z-6Z, vi-8-63 (R. L. Langston, CISX
SONOMA CO.: Cloverdale, Zo'cf, I9, iv-15-56 (R. P. Allen,
CAS, CIS); Eldredge, Zd'd',Z$$, v-16-17, iv-Z-17 (J. A. Kusche,
LACM); Healdsburg, Id, iii-Z5-53 (R. P. Allen, CIS); Jenner,
ld,vii-Z-59 (R. L. Langston, CIS); Z mi. S. Jenner, I9, iv-Z4-
60 (P. A. Opler, PAO) ; meadows east of Mark West Springs,
Id, v-9-60 (N. La Due, NLD); Petaluma, Id, I9 , V-Z6-49 (R. P.
Allen, CIS); Petrified Forest, 1$, iv-Z5-6l (La Due, NJ.D);
7{2) : 65-H6, WfiH
NEARCTIC EUCHLOE
75
Russian River, Zd'd', iii-Z3-33 ( J. A. Comstock, LACM),
MENDOCINO CO.; hills west of Boonville, 1$, vi-1-57 (J. A.
Powell, CIS). TULARE CO.; South Fork Kern River, nr. Deer
Mountain, Ic^, v-31-47(C. Smith, CIS). TUOLUMNE CO. :
Locality unknown. Id*, vii-Zl-30 (Bohart, CAS); Sonora Pass, 9600',
Id*, vii-5-65 (N. La Due, NLD); Sonora Pass, Deadman Creek,
3(3* d*, 1$, vii-3-59 (P. A. Opler, PAO); Tioga Pass, Yosemite
N. P., 2d‘d,29$, vii-13/16-34 (AMNH). YOLOCO. ;. Knight°s
Landing, I9, v- ? - 54 (N. La Due, NLD); 10 mi. W. Winters,
Putah Ck. , I9, v-24-66 (P. Opler, PAO); Yolo Bypass, nr.
Eryte, ZcfcT, lv-6-48. Id”, iv-ll=48 (C. D. MacNeill, CIS); Yolo
Bypass, nr. Davis, Id*, iv-20-30 (C. W. Herr, AMNH); Priest R. , Id*,
v-9-27 (AMNH), ZcTd*, v-10/l9- 26 (C. W. Herr, AMNH). CARIBOU
CO.; SodaSpgs., Id*, 1$, vi-23/24-33 (G. H. and J. L. Sperry,
AMNH). SHOSHONE CO.: Wallace, Id*, v-17-25 (AMNH).
MINNESOTA. ST. LOUIS CO. ; Ash River Trail, Id, vi-17-67
(J. H. Masters, POA), MONTANA. GLACIER CO. : Ptarmigan
Tunnell, Glacier N. P. , 7200', Id*, vii- 26- 64 ( J. G. Edwards,
JGE); Swiftwater L. , Many Glacier Chalet, Glacier N. P, , 1$ ,
vii-4-30 (E. C. Van Dyke, CAS). NEVADA. ELKO CO. : vie.
Arthur, Id*, vi-20/30-29 (E. Schiffel, AMNH); Jarbidge Mts. ,
Id", 1$, vii-12-64 (J. Lane, JL); Wells, Id*, v-23-54 (M. Cazier,
AMNH), WHITE PINE CO. ; Mt. Wheeler, 17d'd', 13$$, v-19/24-29,
v-30-29, vi-2/6-29, vi-8/l0-29 (F.W. Morand, AMNH), OREGON.
BAKER CO.: Baker, 3300', IcT, v-27-57 (J. H. Baker, AMNH);
Pine Ck. , 4100', 1$ vii- 7- 57 (tilden, JWT). HARNEY CO.; Devine
Cyn. , Hwy. 395, 12 mi. NNE Burns, 4800', Id*, vi-1-65 (R. L.
Langston, RLL). JACKSON CO.: Siskiyou Summit, 4522', Id”,
vii-2-58 (R. L. Langston, CIS). JOSEPHINE CO.: Green Mtn.
Spgs. Summit, 1$, vi-10-64 (Tilden, JWT). MALHEUR CO. ;
Ontario, Huntington Rd. , Id*, iv-30-41 (CAS). WALLOWA CO.;
Chief Joseph Mtn. , 2$$, vii-4-52, vii-8-52 (Sperry, AMNH);
Joseph, 4500', 2$$, vii-?-54 (N. Crickmer, AMNH); Wallowa
L, , 4500', Id, 1$, vi-14-39 (CAS), SOUTH DAKOTA. LAWRENCE
CO.: vie. Clayton Draw, 6500', 6dd, vi-17-67 (J. Nordin, PAO);
Spearfish cyn. , 3dd, 2$$, vi-26-39, vii-1- ? (AMNH). UTAH.
CACHE CO.: Logan, Id, vi-12- 33 (AMNH). DAVIS CO.; Mueller Park,
5600', Wasatch Mts. , 6dd, 2$$, v-16-63, v-18-63 (K. Tidwell,'
OAS); Mueller Park, 5400', Wasatch Mts., Id, v-2-63 (K. Tidwell,
OAS). ELDER CO.; nr grigham. Id, 2$$, vii-7/8-25 (LACM).
EMERY CO.: Mohrland, Id, vi-23-? (O.S. Johnson, AMNH),
JUAB CO.; Eureka, Id, v-23-20 (T. Spalding, AMNH). SALT
LAKE CO. : Salt Lake City, 2dd, iv-18-43 (T.B. Ziegler, AMNH);
Salt Lake City, City Creek, 6dd, 1$, iv-22-40, v-11-30, vi-13-44
(CAS, LACM). SAN JUAN CO.; La Sal Mts. , Burro Pass, 10, 500',
Id, vii-22-36 (AMNH); La sal Mts., Gold Hill, 10, 600', Id, 1$,
vii-11-33 (A. G, Richards, jr. , AMNH). SEVIER CO.: Fish L. ,
Id, vii-17-49 (Gertsch, AMNH). TOOELE CO.: 13 mi. SW
Grants ville. Loop Camp, 7400', Id, vii- 3- 60 (R.uv.ige, AMNH).
UTAH CO.; Dividend, Id, v-7-? (T. Spalding, AMNH); Payson
76
OPLER
J. Res. Lepid.
Cyn. , vi-19-33, vi-22-33, vii-7-33 (AMNH); Provo, Slate
Cyn. , Icf, v-14-19 (Spalding, AMNH); Vineyard, 2d'd', 2$$,
V- 22/24- 22 (T. Spalding, AMNH)„ WASHINGTON. OKANAGAN
CO. : Alta L. , 2d'd', iv-29-51(A. Anderson, RLL), YAKIMA
CO]: Bear Cyn. , 2800', 2d'd' (E. J. Newcomer, AMNH);
Bear Cyn., 3200', Id, vi-17-62 (E, J. Newcomer, AMNH);
Little Naches R. , 1$, v-29-59 (E. J. Newcomer, AMNH); Mill
Ck. , 1400', Id, iii-31-62 (E. J, Newcomer, AMNH); Mill Ck. ,
1500', 1$, v-16-62 (E. J. Newcomer, AMNH); Mill Ck. ,
1800', Id, iv-11-62 (E. J. Newcomer, AMNH); Oak Ck. ,
3000', 1$, vi-13-62 (E. J. Newcomer, AMNH); Wenas Ck. ,
1800', 1$, v-14-60 (E. J. Newcomer, AMNH); Wenas Ck. ,
2000', 4dd, iv-26-61 (E. J. Newcomer, AMNH). WYOMING.
TETON CO. : Grand Teton N. P. , 6700', I?, vi-30-53 (J. A.
Ebner, RLL); Jackson Hole, Id, vi-I7-49 (CAS); Jackson
Hole, Moose P. O. , 2dd, v-20-24, v- 23- 24 ( A. B. Klots, AMNH);
base of Teton Range, 1$, vii-11-25 (LACM); Togwottee Pass,
2dd, vi-21-41 (CAS); Yellowstone N. P. , Dunraven Pass, Id,
Id, vi-25-30 (VanDyke, CAS); Yellowstone N. P. , Roosevelt
Camp, 4^9, vii-26-30 (E. C. VanDyke, CAS); Yellowstone N. P. ,
Roosevelt Lodge, I9, vii-3-38 (E. C, Van Dyke, CAS); Yellowstone
N. P„ , Tower Falls Rd. , 1$, vi-7-37 (CAS); Yellowstone N. P. ,
west entrance, I9, vi-14-30 (E. C. VanDyke, CAS). SUBLETTE
CO.: Willow Ck. Rgr. Sta. , 8000- 8900', Id, vii-14- 39 (Klots,
AMNH).
Euchloe ausonides coloradenses (Hy. Edwards
COLORADO, county unknown: 4dd (AMNH), Id, vii-25-55
(Renk, PAO), Id (T.L. Mead, AMNH). BOULDER CO.: Boulder,
2dd, vi-8-22, vii-5-02 (AMNH, LACM); Four Mile Cyn., Id,
vi- 28-52 (D. Eff, JWT), Id, 2$$, v-6-62, v-28-61 (D. Eff,
OAS); Gregory Cyn., I9 , v-19-64 (D. Eff, OAS); Lefthand Cyn.,
v-30-57, vi-8-57 (D. Eff, JWT), Id, v-25-57 (D. Eff, Oas);
Magnolia Rd. , 2$$, v-30-54 (D. Eff, JWT); Mesa Trail, Boulder,
Id, v-12-54 (D. Eff, JWT); Packer's Gulch, 1$, vi-8-57 (D. Eff,
JWT), 3dd, 1$, v-6-62, v-16-63, v-28-61 (D. Eff, OAS); Sunshine
Cyn., 2dd, I9 , iv-27-61, v-5-62 (D. Eff, OAS). CLEAR CREEK
CO.: Beaver Ck. , Hwy. U. S. 40, Id, vi-18-60 (J. Scott, OAS).
3dd, vii-6-61 (Rindge, AMNH). DOLORES CO. : Rico, Id,
vii- ?-? (Oslar, AMNH); Rock Ck. , vie Colorado Springs,
8500-8700', 6dd, vi-5-38, vi-26-41, vi- 30/vii- 4- 39, vii-4-38
(A. B. Klots, AMNH). GILPIN CO.: Hideaway Park, 8715', 1$,
vi-23-54 (L. Martin, LACM). GUNNISON CO. Almont, 2dd, 1$,
vi-20/30-25 (AMNH); 10-20 mi. SE Crented Butte, Cement Ck. ,
9800-11, 000', Id, 1$, viii-8-6l (Rindge, AMNH). JEFFERSON CO,;
Clear Ck. Cyn., I9, v-26-63 (D. Eff, OAS); Golden, Chimney
Gulch, Id, iv-20-04 (Oslar, AMNH); Indian Hills, Id, v-14-38
(AMNH); Mother Cabrini Shrine, 1?, vi-18-60 (J. Scott, OAS).
LARIMER CO.; Estes Park, 2dd, 1$, vi-10-44, vi-16-44
(CAS); Long's Peak, 2dd, vi-15-22 (AMNH); Red Feather
7(2) : 65-H6, 1968
NEARCTIC EUCHLOE
77
Lakes, 8000', 4c/d', vi-19-29 (A. B. Klots, AMNH); Rocky Mtn,
N. P. , Scfd*, vi-?-31 (J. L. Sperry, CAS, LACM), IcT, viii-15-37
(A.B. Klots, AMNH), Id*, 2??, vii-3-35 (L. Hulbirt, JWT).
MINERAL CO,: locality unknown, Icf, vii-1- 40(AMNH), PARK
CO, r ElkCk. , 8500', Id*, vi-4-60 (J. Scott, OAS); Hall Vy. ,
Id*, vi-15-02 (LACM); Hall Vy. , 10, OO'-IO, 500', 4d'd', 1$,
vii-2/3-41, vii-13/15-35 (A, B. Klots, AMNH); South Park, Id*,
vii-29-04 (Oslar, AMNH). ROUTT CO.: Steamboat Springs,
6000', Icf, vii-4-50 (Lot, CAS). SAN MIGUEL CO.: Ophir,
Icf, vii~?-14 (AMNH); San Miguel, Icf, (Oslar, AMNH);
Telluride, Icf, vi-15-04 (LACM); Telluride, Cornet Ck. ,
11, 000*, Id*, vii-9-19 (AMNH). SUMMIT CO.: Fremont Pass,
11,316', Id*, 1$, vi-22- 54 (Martin, LACM). TELLER CO. ;
Green Mtn. Falls, 10,000', 2d'd', vii-l/7-? (AMNH). . NEW
MEXIGO. county unknown; Aspen Ranch, Sangre de Cristo
Range, 9000', Id, vi- 30- 35 (A. B. Klots, AMNH). BERNALILLO
CO.: San Antonio, Id*, vii-9-36 (R. Kaiser, AMNH). SAN
MIGUEL GO.; Cowles, Windsor Cyn. , 8000-8500*, IcT, 1$,
vii-3-35 (A.B. Klots, AMNH). WYOMING. ALBANY CO.:
Centennial, Univ. Wyoming Camp, 9600', 9d'd', 8$$, vi- 26/vii- 6- 29
(A.B. Klots, AMNH, CIS); Woods Landing, Id*, vi-18-55
(G. DeFoliart, JWT).
Euchloe ausonides mayi Ghermock and Chermock
CANADA. - MANITOBA. Beulah, 2dd', vi-20-37 (AMNH);
Duck Mountain Provincial Park, Id*, Id*, vii-1-36 (J. S. Nordin,
PAO); Duck Mountain Provincial Park, vie. Ketchum Hill,
2d'd', vii-15-67 (J. H. Masters, PAO); Herchman, mi, 412,
2d'd, vi-23-32, vi-24-32 (AMNH); McCreaft, Id, vi-22-38
(AMNH); Riding Mts. , 2dd, vi-18-37, vi- 29- 34 (F. H. and R. L.
Chermock, AMNH) PARATYPES, 1$, vi-8-36 (AMNH), 2$$,
vi~27-32 (J. F. May, LACM); Riding Mts., Kelwood, 1$,
vi-12-31 (CAS), Id, 1$, vi-2/26-29 (J. F. May, AMNH); Riding
Mts., Trail to Grey Owl's Gabin, 6dd, vii-24-67 (J. H. Masters,
PAO); Riding Mountain National Park, I5, vi-19-60 (J. F. May, AMNH)
AMNH); Sand Ridge, Id, vi-11- ? (AMNH) PARATYPE, 2dd,
vi-26-36 (AMNH).
Euchloe creusa creusa (Doubleday)
CANADA. - ALBERTA. Banff, Id, vi- 21- 25 (G. P. Englehardt,
AMNH), Id, v-29-22 (C. B. D. Garrett, LACM); vie Banff, 2dd,
vi-21/31-29 (J; F. May, AMNH); Banff, Cascade Mtn. Amphi-
theatre, 7000', 1$, vi-29-25(0. Bryant, AMNH); Banff, Rundle
Mtn,, E face, 5000'-7000', Id, vi-25-25 (O. Bryant, AMNH);
Bow Lake, 2dd, vi-7-23 (G. C. Hall, AMNH); Lagg an. Id,
78
OPLER
/. Res. Lepid.
vii-1-17 (AMNH); Nordegg^ 1$^ vii-1-66 (J. Lane, JL); Atlin, 6d'd',
vi-7-?, vi-20-?, vi-21-?, vi-28-? (AMNH)„ YUKON TERRITORY.
Whitehorse, Icf, vi-9-23 (LACM), Icf, 1$, vi-9-66 (J. A. Ebner,
PAO).
UNITED STATES, - ALASKA. Eagle, Id*, vi-8~36 (F,
Grinnell, LACM); Klutina Lake, 2d'd', v-15-50 (W. C, Frowne,,
DLB),
Euchloe creusa, Saskatchewan segregate
CANADA. - SASKATCHEWAN. Prince Albert, Nesbit
Provincial Forest, ScTcf, 3$$, v-27-51, vi-2-51 (Aschim, AMNH),
Euchloe hyantis hyantis (Edwards)
CALIFORNIA. HUMBOLDT CO. : locality unknown, Icf, no
date (Hy. Edwards, AMNH), 1$ (Koebele, CAS). LAKE CO. : 9 mi.
N Upper Lake, Icf, iv-4-62 (Langston, RLL). MENDOCINO CO.:
Hopland, 2$$, v-14-32 (CIS). SHASTA CO.: Redding, Id, 1$,
iii- 2-63, iii-12-63 (D. L. Bauer, DLB). SISKIYOU CO.: Castle
Lake, 2cfcf, 1$, vii-12-58 (J. Powell, CJS); Lake Mountain, Icf,
vii-4-63 (P. A. Opler, PAO); nr. Mt. Shasta, 1$, vi-10~41 (CIS);
Selad, 1$, v-8-59 (R. P. Allen, CAS); n mi, N Mt. Shasta City, 2cfcf,
1$, vi-25-58 (J. Powell, CIS), Icf, vii-3-58 (N. La Due, NLD).
SONOMA CO.: locality unknown, Icf, iv-17-38 (LACM); Cazadero,
Icf, 1$, iv-17-30 (CAS, CIS), Id*, 1$, v-3-31(W. C. Wood, AMNH),
2cfcf, I9, v-19-29 (Bohart, CAS); Cloverdale, Icf, iv-25-57 (R. P,
Allen, CAS); 2 mi. E Du ncan Mills, Icf, I9 , vi-1-57 (R. L,
Langston, RLL); Geysers, Icf, v-10-38 (E. C. Johnston, AMNH), Icf,
iv- 8-61 (Tilden, JWT), Icf, iv-22-59 (M. Doudoroff, CIS); 3 mi.
NE Guerneville, 2d'd', iv-8-55 (Langston, RLL). TRINITY CO.:
Trinity Center Delta, Trinity Mts. , 1$, v-22-25 (R. F. Sternitzky,
AMNH).
Euchloe hyantis. Sierra Nevada segregates
CALIFORNIA, county unknown: Locality unknown, acc. no.
3576, Icf, no date (Hy. Edwards, AMNH); Feather River, 7cfcf,
599, vii-4/18-28 (AMNH), 2cfd', iii- 25- 32 (AMNH); Lake Tahoe,
Id*, vi-11-35 (AMNH), Id, vi-13-30 (R. F. Sternitzky, CAS), Id,
v- 29-30 (LACM); Sierra Nevada, acc. nq,4278. Id (Hy. Edwards,
AMNH); Yosemite, acc. no. 4278, 2dd (Osten Sacken, AMNH);
Yosemite N. P. , Id, v-13-27 (CAS); Yuba River, I9, v-29-27 (CAS).
ALPINE CO. : Hope Vy. , 5dd, vi-9-48 (J. W. MacSwain, CIS).
AMADOR CO.: peaks over Carson Spur, 8200', 4dd, vi-20-61
(N. La Due, NLD); rocks over silver Lake, 8000*, 2dd, I9,
vi- 19-61, vii-2-62 (N. La Due, NLD). CALAVERAS CO.:
Camp Wolfeboro, B. S. A. , N. Fk. Stanislaus River, 5600', 299,
vi-18~54, vi-21-54 (P. A. Opler, CAS, PAO); Dorrington, 2dd,
iv-12-33 (CIS), 2dd, 19, vi-10-30 (AMNH); EL DORADO CO.:
Echo Lake, Id, vii-22-63 (N. La Due, NLD); nr. Echo Lake,
7(2) : 65-86, 1968
NEARCTIC EUCHLOE
79
ZcfcT, 1$, vii-6-40, vii-9-40 (CIS); Mt. Tallac, 9 785', Id*, vii-Z7-39
(F, H. Rindge, AMNH); Wright's Lake, Id, vii-2-48 (J. W. MacSwain,
CIS). MARIPOSA CO. : w mi. n Bear Vy. , Hwy. 49, Id, iv-17=-6l
(Opler, PAO), Id, 1$, iv-26-65 (R. E. Stanford, Hughes-Stanford
Coll'n); Boundary Hill, Yosemite, Id, vii-10-54 (R. P. Allen, CIS);
El Portal, 3dd, 1$, iv-lL64 (Opler, PAO), 2??, iv-27-21, vii-2-21
Stanford-Highes Coll'n); Glacier Pt. , 2dd, vi-28-21, vii-2-21
(J. A. Comstock, LACM); above Indian Flat, 21dd, 7$$, iv-14-63
(La Due, NLD); 1/ 2 mi. E May Lake, Yosemite N. P, , 8900',
Id, vii-4-46 (F. H. Rindge, AMNH); Tamarack Flat, Y. N. P. ,
Id, vii-3-54 (Tilden, JWT); Wawona, 2dd, v-25-? (L. B. Woodruff,
AMNH); Yosemite Vy. , 3880-4000', 1$, vi-l-38(R. M. Bohart, CAS),
Id, vi-3-38 (N. Hardman, AMNH). NEVADA CO. t Donner Pass, Id,
vii- 4-62 (J. Powell, CIS). PLACER CO.: locality uninown, I5 (AMNH),
3dd, 259, "June" (Koebele, CAS); nr. Auburn, N. Fk. American
River, 800', 20dd, 49$, iv-18-61, iv-21-63, iv-22-63, iv-23-62,
v-1-67 (La Due, NLD); Mt. Judah, north slope, 2dd, vii-16-63,
viii- 6-63 (La Due, NLD). PLUMAS CO.: Feather River, Id,
iii- 25-30 (R. Wind, JWT). SIERRA CO. : Bassets, I9, vii-8-67
(Opler, PAO); Gold Lake, 2dd, 899, vii-14-21, vii-15- 25, vii-16- 21
(J. D. Gunder, AMNH), 2dd, vii-1-35, vii-10-35 (L. M. Martin,
LACM); Id, vii-2-63 (La Due, NLD), Id, vii-19-67 (O. Shields, O
OAS), Id, vii-8-67 (Opler, PAO); 3 mi. E Gold Lake, Id, vii-1-61
(D. Dirks, OAS); Salmon Lake, 2dd, vii-27-? (CAS); Shenanigan
Flat, 2dd, vi-17-67 (Opler, PAO); Yuba Pass, I9, vi-28-60
(Tilden, JWT). TULARE CO. : Sequoia N. P. , Id, iv-10-30 (CAS).
TUOLUMNE CO. ; Hog Ranch, 4600', 1 mi. E Mather, I9, vi-23-64
(O. Shields, OAS); nr. Sonora Pass, 5dd, 299, vi-20-47 (C. D.
MacNeill, CIS); Strawberry Lake, I9, vi-10-32 (CAS); Tuolumne
River, nr. Tuolumne City, Id, iv-17-61 (P, A. Opler, PAO), Id,
iv- 27-65 (R. E. Stanford, H"i:ghes- Stanford Coll'n). NEVADA.
DOUGLAS CO.: Kingsbury Grade, I9, vi~18-67 (Opler, PAO).
Euchloe hyantis, Mt, Pinos Block segregate
CALIFORNIA. KERN CO. : Lebec, Id, vii-8-56 (Opler, PAO);
Sand Springs, 6000', Id, v-28-39 (Rindge, AMNH). VENTURA CO. :
Chuchupate Rgr. Sta. , 3dd, 299, v-1-59, v-5-59 (J. Powell, CIS),
Id, 299, v-8-59 (P. D . Hurd, CIS), 4dd, v-14-60 (Spier, PAO); Cuyama, 1
Cuyama, Id, iv-8-58 (R. P. Allen, CIS).
Euchloe hyantis. Peninsular Ranges segregate
CALIFORNIA. RIVERSIDE CO.: Palm Springs, Id, iv-3-25
(E. P. Van Dyke, CAS); Pinyon Flat, Id, iv-20-62 (MacNeill,
Rentz, and Brown, CAS), 3dd, iv-13-63 ( Langston, RLL), 2dd,
Id, iv-7-63 (Langston, CIS), Id, iv-12-63 (G. Tamaki, CIS);
80
OPLER
/. Res. Lepid.
Santa Rosa Summit^ 7 mi. E Anza, Icf, iv-18-62 (NacNeill,
Rentz, and Brown, CAS). SAN DIEGO CO.: Agua Caliente, 4d'd',
iv-6-50, iv-6-52, iv-14-51, iv-25-53 (Powell, CIS); 2 mi.
SE Banner, Id, iii-31-63 (O. Shields, AOS); 3 mi. E Banner,
2500', 2d'd', i-21-58, ii-9-58 (F. Thorne, PAO); Buckman Spring,
Id, iii-14-63 (Powell, CIS); Descanso Rgr. Sta. , Id, iii-30-6l
(Langston, RLL); Horse Haven Gorge, Laguna Mts. , Id, v-1-65
(O. Shields, AOS); Jacumba, Id, iv-9-52 (Powell, CIS); 3 mi.
NW Jacumba, Id, 1$, iii-31-63 (F. Thorne, PAO); 4 mi. E
Jacumba, 3200', Id, iii-27-66 (O. Shields, AOS); Laguna Mtn,
Recr. Area, 4dd, 1$, iv-22-51, v-11- 51 (Langston, RLL);
Monument Peak, Laguna Mts., 3dd, v-11-52 (Langston, RLL);
nr. Monument Peak, Id, iv-22-51 (Langston, RLL); Mt Laguna,
Id, vi-21-63 (P. Welles, CIS); Pine Vy. , 3600', Id, v-1-65
(O. Shields, AOS); Scove Cyn. , 3mi. N Mt. Laguna Jet., Id,
iii-27-61 (Langston, CIS); Storm Cyn., Laguna Mts., 5500', 9dd,
1$, iv-21/24-62 (F. Thorne, LACM, PAO).
Euchloe hyantis andrewsi Martin
CALIFORNIA. SAN BERNARDINO CO.: Cedar Pines, Id,
v- 13-28 (LACM); Crestline, 5000', 14dd, 2$$, v-26-40,
vi- 7-39, vi-10-49 (Rindge, AMNH), 5dd, 1? (W. T. Meyer,
LACM, CIS, JWT); Crestview, 3dd, 1?, v-24-42 (Rindge, AMNH);
Fish Camp, Id, vi-18-36 (C. M. Dammers, LACM); nr Lake
Arrowhead, Id, v-25-47 (C. Smith, CIS); nr. Lake Arrowhead,
Crestline Hwy. , 3dd, 2$$, v-19-35, 1?, vi-15-36 PARATYPES,
7dd, 2$9, vi-8-?.6 TOPOTYPES (R. H. Andrews, LACM, JWT);
Running Spring, 2dd, iv-2-40 (C. Smith, CIS), Id, v-25-47
(LACM); Sugarloaf Mtn., 8000', vie. Big Bear Lake, Id,
vi-19-66 (J. Lane, JL).
Euchloe hyantis lotta Beutenmuller
CANADA. - BRITISH COLUMBIA. Cranbrook, 6dd, 5$$,
iv-18-13, iv-19-13, iv-21-13, iv-27-13, v-2-10, v-5-10, v-30-11
(C. Garrett, AMNH); Lilooet, 2dd, iv-20-18, iv-28-17 (AMNH);
Oliver, 2$?, iv-24/25-23 (AMNH).
UNITED STATES. - ARIZONA. GILA CO. : Globe Id, iii-30-41
(AMNH). MARICOPA CO.: Apache Lake, Id, iii-7-53 (G. W.
Kirkwood, LACM). MOHAVE CO.: Peach Springs, 2dd, iv-12-40
(Rindge, AMNH). PIMA CO.: Brown Cyn,, Baboquivari Mts., 1$,
iii- 31-53 (L. M. Martin, LACM); Catalina Mts., Id (AMNH);
Fresnal Cyn. , Baboquivary Mts. , 3dd, Iq, iii-14-59 (K. Roever,
JWT); Madera Cyn, , Id, iii-28-6l(0. Shields, AOS), 4dd, 1$,
iv- 2/3-53, iv-15-54 (L. M. Martin, LACM). SANTA CRUZ CO. :
Patagonia, 2dd, iii-24-38 (CIS, JWT). YAVAPAI CO.:
7(2) : (i5-H6, n)(iH
NEARCTIC EUCHLOE
81
Mingus Mtn. , Id-, iii-31-50 (D, L. Bauer, LACM). CALIFORNIA.
INYO CO.: Aguerreberry Camp, 53Z0', Scfd*, I5 , iv-8-60,
iv-20-58 (Langston, RLL); Alabama Hills, 6 mi. W Lone Pine,
8d'd', 1$, iv-9-60 (Langston, RLL); Alabama Hills, Tuttle Cyn. ,
3 mi. W Lone Pine, 1$, iv-9-60 (Langston, RLL); Argus Mts. ,
Zd'd', 145$, iv-?-1891, v-7-1891 (CAS); summit Argus Mts. , Hwy. 190,
Zd'd', iv-13-60 (Opler, PAO); nr. Haiwee Summit, 1$, iv-1-47
(C. Smith, CIS); 5 mi. N Little Lake, I9, iv-10-62 (J. W. MacSwain,
CIS); 9 m i. W Lone Pine, Icf, iv-9-60 (Langston, RLL); Panamint
Mts., Id*, iv- 7-1891 (CAS); 3 mi. E Skidoo, Panamint Mts., 5500*.
Icf, iv-13-57 (Langston, RLL); Surprise Cyn,, Panamint Mts., Icf,
iv-24-57 (J. Powell, CIS); Wild Rose Springs, Panamint Mts.,
3500', Icf, 1$, iv-12-57 (Langston, RLL), KERN CO. : locality
unknown, 1$, iv-15-27 (J. S. Garth, AMNH); Mojave, 3cfcf,
iv-15/17-38 (AMNH); 22 mi, E Mojave, Icf, iv-1-26 (AMNH);
8 mi. W Mojave, 2cfcf, iv-11-58 (J. W. MacSwain, CIS); Red Cyn.,
Mojave Desert, Icf, iv-18-30 (CAS); 9 mi. N Ricardo, east branch
Last Chance Cyn. , Hwy. 6, Icf, iv-10-60 ( C. A. Toschi, CIS); 5 mi.
E Roamond, Icf, iii-27-60 (K. Shea, PAO); Taft, 1$, iv-?-? (AMNH)
Walker Pass, 5200', 1$, vi-17-67 (F. Thorne, PAO). LASSENCO :
1 mi. W Hallelujah Jet., 2cfcf, vi-17-67 (P. Opler, PAO). LOS
ANGELES CO.: locality unknown, Icf (CAS), Icf, v- ? - ? (CAS);
Boquet Cyn., Sierra Madre Mts., 1$, iv-12-25 (LACM); Desert
Spgs. , 2cfcf, v-7-63 (Langston, CIS); Littlerock, llcfcf, 5$$,
iii- 17/25-28, iii-21/22-31, iii- 2l/iv- 2- 32, iii-25-33 (AMNH, CAS,
LACM), 2cfcf, 3$$, iii-16-35, iv-12-36 (G. Heid, GAS), 3cfcf, 3?$,
iv- 6-39 (C. Smith, GIS); 2d'cf, iv-4-32 (R. W. L. Potts, CIS), Icf,
iv-?-36 (J. Fischer, RLL), Icf, iv- 20- 50 (C. D. MacNeill, CIS),
35cfcf, 12$$, iii-28-64, iv-3/ll-55, iv-9-60, iv-ll/l2-54 (P. Opler,'
PAO), 2cfd', iii-25-59 (Tilden, JWT); nr. Littlerock, Zefcf, iii-16-40
(C. Smith, CIS); nr. Littlerock Dam, Icf, iii- 30-41 (CIS); Llano,
3000', Icf, iii-11-34 (LACM), 1$, ii-15- 37 (J. A. Comstock, LACM);
Lovejoy Buttes, Mojave Deseit, 2cfcf, iii-18-47, iv-6-41 (C. Smith,
CIS); Mint Cyn., +cfd', 2$$, iv-18-34, iv-20-28 (AMNH, LACM);
Palmdale, 3cfcf, iv-5-30, v-1-37 (AMNH), 1$, iv-14-27 (J. S.
Garth, CIS), Icf, iii-21-47 (LACM); Switzers, wcfcf, 1$, iii-21-32
( W. A. Evans, CAS); Valyermo, 1$, iv-3-38 (AMNH), 6cfcf,
iii-28-37, iv-28-35 (G. Heid, CAS). MODOC CO.: 6 mi. W
Alturas, 2cfcf, 1$, v-23-56 (Langston, RLL); Cedarville, 1$,
vi-4-35 (E. C. Johnston, AMNH). MONO CO. : Mono Lake, Icf, vi-17-19
(AMNH). RIVERSIDE CO. : 1 mi. N Desert Center, 1$, iv-11-58
(W. E. Ferguson, CIS); Split Rock Tank, Icf, iii- 22- 3 9 (AMNH).
SAN BERNARDINO CO. : Adelanto, Icf, iii- 20- 31 (C. M. Dammers,
LACM); Baldy Mesa, 1$, iv-9-37(J. A. Comstock, LACM); Barstow,
82
OPLER
/. Res. Lepid.
\cf , iv-8-31 (CAS); above Bonanza King Mine, Providence Mts, ,
Zcfc/, iv-7-66 (P. Opler, PAO), nc/d, 7$?, iii~15-67 (G,
Gorelick, GG); 16 mi. SW Cima, 5000*, Odd*, 2^$, iv-2-66
(P. Opler. PAO); 1/2 mi. W Cotton wood Spring, 4000-5000',
Granite Mts., nr. Essex, 4^^, iii-Z2-67 (O. Shields. AOS);
Kramer, Id*, iv-2-32 (AMNH); Mitchell Caverns, Providence
Mts., Icf, iv-10-52 (F. Thorne, FT); ridge just west of Mitchell
Caverns State Park, Id*, iv-14-65 (O. Shields, A. OS); Paradise
Vy. , Zcfd, 295 (CAS); Phelan, 5cfd', 5$$, iv-10~38, iv-11-37,
iv-15-33, iv-18-50 (AMNH, LACM), 3d'd, 2$?, iii-28-35,
iv-4-37, iv-13-35, iv-14-37 (F. Estes, AMNH), 5d'd, 1$, iii-20-31,
iv- 11-30, iv-18-30 (C. M. Dammers, LACM); 2 mi. S Phelan, 1$,
v- 7-63 (CIS); Quail Springs, Id*, iv-15-38 (AMNH); between
Randsburg and Kramer, lOcfd*, 4$5, v-5-27 (T. Craig, CAS);
San Bernardino, 1$, iii-15-14 (V. L. Clemence, LACM); Sheep Ck. ,
Sd-d, 5$$, iv-22-28, iv-24-29 (LACM), 5dd, iii- 20- 31 (C. M,
Dammers, LACM); Victorville, Zdcf, I9, iv-18-31 (CAS), Icf,
1$ , iv-15-57 (Tilden, JWT); Upper Volta, Phelan, 1$, iii-6-33
(J. L. Sperry, CAS); Yucca Vy. , 25 mi. W 29 Palms, Icf, iv-8-55
(F. Thorne, FT). COLORADO. MESA CO. : Black Ridge, Icf,
v-lO-46 (AMNH), Icf, v-17-6l (D. Eff, AOS); Black ridge Breaks,
Frita, Icf, vi-5-44 (CAS); Black Ridge, Coal Mine Point, 3cfcf,
v- 11-63 (O. Shields, AOS), Zd-d*, v-17/18-61 (D. Eff, AOS);
Decil's Cyn. , 5000', 2$$, iv-23-40 (AMNH); Glande Park, 7000*,
5cfcf, 2$$, iv- 20-40 (AMNH). MONTEZUMA CO.: Mesa Verde,
Icf, iv-21-40 (LACM). IDAHO. ADA CO. ; Kuna, ecfcf, S?? (AMNH).
BINGHAM CO. : Blackfoot, Zcfcf, iv-18- ? (AMNH). BONNER CO.;
Priest River, Icf, vii-9-20 (AMNH). NEVADA, county unknown:
Icf (AMNH). ORMSBY CO.: Eagle Valley, Carson City, 4700',
1$, v-10-61 (P. Herlan, NSM). WASHOE CO. ; Reno, Icf, v-9-18
(LACM), 19, v-7-08 (F. Burns, LACM); hills 2 mi. N reno. Id”,
vi- 18-67 (P. Opler, PAO). WHITE PINE CO. : Mt. Wheeler, 4cfd',
v-19/24-29 (F. W. Morand, AMNH). NEW MEXICO. HILDALGO CO. :
Rodeo, Id”, 1$, iii-9-38 (Tilden, JWT). RIO ARRIBA CO. : I mi, '
E Capulin, llcfd”, 1$, iv-11-63 (J. Scott, AOS); Em budo. Id”, iv-2I-62
(J. Scott, AOS). OREGON. BAKER CO.: Baker, Tcfd”, v-20-57,
v-21-58 (J. H. Baker, AMNH). MALHEUR CO.; Huntington Rd. ,
Ontario, I9, iv-30-41 (CAS); nr. Rockville, Icf, v-18-61 (J. Baker,
AOS). UTAH. BEAVER CO.: Beaver Ck. Hills, 3cfcf, v-?-?,
v-?-? (AMNH). JUAB CO.: Eureka, 9d'd”, 1$, v-7/21-11 (T.
Spalding, AMNH), Id”, 29$, v-29-20 (LACM). SALT LAKE CO. :
Salt Lake City, I9 , v-?-? (AMNH). TOOELE CO.: Stockton,
Sd-d”, 399, iv-29-07, v-6-28, v-11-09, v-30-07, vi-8-06 (T.
Spalding, AMNH, JWT); North Willow Ck. , Stansbury Mts. ,
5400', Jdd”, v-15-63 (K. Tidwell, AOS). WASHINGTON CO. :
ZionN. P. , Id”, iv-18-27 (CAS), WAXHINGTON. BENTON CO. :
Prosser, Icf, iv-25-21 (W, Lord, LACM); 10 mi. NW Richland,
7(2) : 65-^6, 196H
NEARCTIC EUCHLOE
83
3d'd', iv-14-6Z (R. E. Woodley, AOS). CHELAN CO.: Port Columbia,
15, iv-16-? (CAS). DOUGLAS CO.: Dyer Hill, 1?, v-7-55 (PAO).
OKANAGAN CO.: Alta Lake, Zcfcf, iv-29-51 (A. Anderson, RLL);
Brewster, Ibd'd', 11$9, iv-20-32, iv-18-54, iv-19-58, iv-28-58,
iv- 30-60, v-1-60 (J. W. Hopfinger, AMNH, JWT); Pateros, Id*, 1?,
v- 2-33 (W. C. Wood, AMNH), Zcf cf , 1$, iv-23-33, v-10-32,
v-15-32 (AOS, CIS). YAKIMA CO. : Priest Rapids, 500', 9d'd',
3$9, iv-2-62, iv-5/8-66 (E. J, Newcomer, AMNH, PAO).
WYOMING. TETON CO. : Jackson Hole, IcT, v-20-24 (AMNH);
Jackson Hole, Moose P, O. , Icf, 2$$, v-16-24, v-20-24, v-23-24
(A. B. Klots, AMNH).
ADDENDA
Euchloe ausonides
CANADA. ~ ALBERTA. Banff, 1 vii-7~02 (J. Fletcher); Calgary,
9 d^, v-29-14 (F.H. Wolley Dod) ; Elkwater, 1 ^ vl-15-29 (J.H. Pepper);
Fort Fitzgerald, 2 dd, 1 vi-30-50 (W.G. Helps); Lethbridge, 2
V-29--29 (J.H. Pepper); Manyberries, 1 vi-4-56 (E.E. Sterns); McMurry,
2 vi-4-53(W. J. Grown); Nordegg, 5 2 vi-13/ 28-21 (J. McDunnough) ;
Waterton Lakes, 1 (?, vi-27-29 (J.H. Pepper), 2 1 vi-20/27-33
(J. McDunnough), 1 <?, vi-14-22 (C.H. Young). BRITISH COLUMBIA. Atlin,
2200* , 13 12 v-28/vi-24-55 (B.A. Gibbard & H. Huckel) ; Clinton,
9 S^, 2 ^o, vi-10-38 (G.S. Walley) ; Fairview, 1 c?, v-6-36 (A.N. Gartrell);
Fort Nelson, 1 d, vi-10-48 (W.R. Mason); Fort St. John, 1 vi-12-27
(P.N. Vroom) ; Garnett Vy. , Summerland, 1 c?, v-13-33 (A.N. Gartrell);
Grand Forks, 1 v-26-37 (J.K. Jacob); Hedley, 6 d'(^, vii-21-33 (C.B.
Garrett); Hedley, Nickel Plate, 5000', 6 1 vii-l6/l7-53 (McGillis
& Martin) ; Hope Mts., 6OOO', 1 d^, viii-2-32 (A.N. Gartrell); Jesmond,
1 vii-17-37 (J.K. JAcob) ; Kamloops, 4 d'J', 3 99, vi-7-37 (J.K. Jacob);
Kamloops, Mt. Lolo, 5 4 QQf v-3l/vi-2“38 (G.S. Walley); Merritt,
84
OPLER
/. Res. Lepid.
3 d'd', v-15-37 (J.K. Jacob) | Midway, 2 d'S', 1 v-Zl-Jl (J.K. Jacob) j
Okanagan Falls, 1 cf, v-15-53 (D.F. Hardwick); Oliver, 1 c^, v-20-59
(R. Madge); Osoyoos, 2000-2500', 3 v-13-53 (Hardwick & McGillis),
6 (?^, 1 v-21-53 (J. Martin), 1 vii-1-53 (McGillis), 1 vi-8-53,
1 (?, vii-3-53 (D.F. Hardwick), 3 4 v-19-38 (J.K. Jacob); Osoyoos,
Anarchist Mt., 2 v-8-36 (A.N. Gartrell); Pavilion Lake, 2 (?{?, 1
vi-6-38 (J.K. Jacob); Seton Lake, 1 c?, vii-29-33 (J. McIXinnough) ; Shingle
Crk., Penticton, 11 8 vi-5-33 (J. McDunnough) ; Victoria, 1 c^,
iv-5-1885, 1 v-26-1882, 1 c?, v-11-04 (No Collector), 1 vi-3-03
(G.W. Taylor). MANITOBA. Gillam, 15 9 vi-l6/vii-7-50 (J.F.
McAlpine) ; Riding Mts., 1 3', vi-9-37 (W. J. Brown), 36 (?c^, 5 (Brown
& J. McIXannough) . NORTHWEST TERRITORIES. Fort Snith, 1 J', v-30-51, 14
10 vi-2/17-50 (J.B. Wallis); Hay River, 1 c?, vi-14-51 (P.R. Ehrlich);
Norman Wells, 7 3*4', 4^^, vi-lO/vil-11-49 (S.D. Hicks), 1 vii-4-49
(W. R.M. Mason); Yelloidcnife, 7 00, 4 00, vi -13/30-49 (E.F. Cashman
& R.R. Hall). ONTARIO. Beardmore, 15 mi. E. , 2 vi-1-58 (P.D. Syme);
Fort Williams, 2 vi-?-64 (No Collector). SASKATCHEWAN. Cut Knife,
Atton's Lake, 4 ^4, 2 v-29-40 (A.R. Brooks); Cypress Hills, 1
Vi -19-39 (A.R. Brooks); Harlan, 1 S', 2 v-26-40, 1 S, v-24-41, 2 SS,
2 0^, V-I2/I4-47 (P.F. Bruggemann) ; Nipawn Nat, For., 1 S, v-25-42
(F.H. Chermock). YUKON TERRITORY. Dawson, I <?, 1 j, vi-IO/21-49 (W.W.
Judd); Dawson, 1500-2000', 11 SS, 12 o^, vi-8/vli-l-49 (P.F, Bruggemann);
Pelly River, below Hook River, 1 o, vii-5-07 (J. Keele); Rampart House,
1 o, vi-8-51 (C.C. Loan); Ross River, 132° 30', 61° 56', 3000', 1 4, 1
vi-21-60 (J.E.H. Martin); Whitehorse, 1 S, vii-4-48 (W.R. Mason), 3 SS,
1 J, vi-1-49 (P.F. Bruggemann), 1 S, vi -10-49 (D.A. Mitchell), 2 p,
Vi -6/21-49 (D.L. Watson), 1 c?, 1 ?-?-21 (E.P. Hawes).
7(2) : 196H
NEARCTIC EUCHLOE
85
UNITED STATES. - ALASKA. Big Delta, 4 v-28/vi-l8-51 (Mason
& McGillis); Moose Pass, Kenai Peninsula, 3 2 vl-20/vii-2--51
(W.J. Brown); Richard Hwy. , Mile 270 , 3 cTcf, vi~4-65 (Mason & McGillis);
Richard Hwy., Mile 275, 3 1 vi“6-51 (Mason & McGillis).
Euchloe creusa
CANADA. - ALBERTA. Banff, 1 J*, vii-7-02 (J. Fletcher), 1 j,
vii-1-07, 1 o, vi-25-08 (F.H. Wolley Dod) ; Banff Nat. Park, 3 2
vli-9-55 (Brown, McGillis & Shewell); Canyon Crk., 4000*, 1 5, vii-1-65
(J.R.H.); Hillcrest, 1 cf, 1 o, vl-20-20 (No Collector); Laggan, 1 d*.
No Date (T.E. Bean); Laggan, Mt. Pinair, 7000', 1 (?, 1 0, vii-17-07
(F.H. Wolley Dod); Moraine Lake, 1 d, vii-20-38 (G.S. Walley) ; Nordegg,
3 56*, 3 o^, vl-lO/22-21 (J. McDunnough) ; Sunwapta Pass, 6600* , Banff-
Jasper Hwy., 1 d*, vii-9--55 (R. Coyles); Waterton Lakes Park, 1 c?,
vii-2-22 (C.H. Young). BRITISH COLUMBIA. Atlin, 2200*, 5 vi-6/26-55
(R.A. Gibbard & H. Huckel) ; Atlin, 4OOO*, 1 (?, vii-26-55 (H.J. Huckel) ;
Golden, 1 v-29-64 (J.R.H.); Kootenay Park, Vemillion River, base of
Mt, Gray, 48OO*, 1 vii-15-55 (R. Coyles); Summit Lake, Mile 392,
Alaska Hwy., 7 ^9, vi-20/vli-8-59 (R.E. Leech & E.E. MacDougall) .
NORTHWEST TERRITORIES. Cameron Bay, Great Bear Lake, 2 SS, 1 9, vii-3-37
(T.N. Freeman); Fort Reliance, Great Slave Lake Region, 1 9, vii-6-24
(J, Russell); Great Slave Lake Region, 1 <5', vi-10-22 (J. Russell); Norman
Wells, 4 1 9? vi-9/l5-49 (W. R.M. Mason); Reindeer Depot, MacKenzie
Delta, 11 00, 3 00, vii-l/12-48 (W.J. Brown & J.R, Vockeroth) ; Saw Mill
Bay, 10 dcf, vl-18/21-48 (D.F. Hardwick); Yello\dcnife, 1 9, vi -18-49
(E.F. Cashman). YUKON TERRITORY. Aklavik, 1 9, vi-l6-53 (C.D. Bird);
86
OPLER
J. Res. Lepid.
Old Crow, 1 o’*, vii-2-51 (C.C. Loan); Rampart House, 8 ^6*, 1 j, v-3l/
vi-31-51 (C.C. Loan & J.E.H. Martin); Sheldon Lake, 131° 37', 62° 54',
3500', 1 o, vii-7-60 (J.E.H. Martin); Whitehorse, 2300', 1 (?, vi-4-49
(P.F. Bruggemann) ,
UNITED STATES. - ALASKA. Anchorage, 2 vi-5-51 (R.S. Bigelow);
Kenai Peninsula, Moose Pass, 2 (?<^, vi-20/ vii-2-51 (W.J. Brown).
Euchloe hyantis
CANADA. - BRITISH COLUMBIA. Garnett Vy. , Summerland, 3 ^cf, 5 ^9,
v-l/5-33 (A.N. Gartrell); Lillooet, 1 v-?-27 (A.W. Phair); Okanagan
Falls, 2000', 3 v-9/15-53 (D.F. Hardwick); Oliver, 7 4 iv-22/
v-7-23 (C.D. Garrett), 3 5 iv-22-33 (A.N. Gartrell), 1 v-16-38
(J.K. Jacob); Osoyoos, 2500', 3 1^, v-13-53 (D.F. Hardwick & J.R. McGillis)
Osoyoos, Anarchist Mt., 1^, v-7-36 (A.N. Gartrell); Penticton, iv-20/
26-31 (A.N. Gartrell); Summerland, 1 iv-20-31 (A.N. Gartrell).
Journal of Research on the Lepidoptera
7(2) : 87-94, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
NOTES AND RECORDS ON
SPECIES IN THE GENERA FOLIA
OCHSENHEIMER AND EUXOA HUBNER
FROM THE WESTERN UNITED STATES
(LEPIDOPTERA: NOCTUIDAE)
JOHN S. BUCKETT
Systematic Entomologist,
Bureau of Entomology
California Department of Agriculture,
Sacramento, Calif. 95814
The majority of the species considered here represent Cali-
fornia records, and “California” has been omitted after the
county when this is the case. One species is also new to Nevada
and where specimens are from another state, the state is indi-
cated. The specimens cited in this paper are contained in the
Bauer-Buckett Collection, Davis; the collection of the California
Department of Agriculture, Sacramento; Entomology collection,
University of California, Davis. In each species, both male and
female genitalic preparations were made.
Polia lilacina (Harvey)
No published records of Polia lilacina (Harvey) have includ-
ed California. Holland (1903) reports distribution in New Mex-
ico, and Crumb (1956) lists Maine, New Hampshire, Connecti-
cut, New York, New Jersey, Pennsylvania, Iowa, North Dakota,
Arizona, Nevada, Utah, Colorado, Oregon, Washington, Mon-
tana and all the adjacent Provinces of Canada. P. lilacina was
orginally described from Brewster, New York.
A specimen was received from Bishop, Inyo County, 3 August
1960 (R.P. Allen) and was found to be conspecific with lilacina.
A male and a female, color form illabefacta (Morrison), were
collected at Fort Dick, Del Norte County, 3 August and 3
September 1962 (J. W. Anderson) respectively. The maculation
of this color form is less clearly defined than in lilacina and was,
for this reason, considered a separate species for some time. The
primaries in this form are suffused with lilac-brown and, thus,
are somewhat obscured although maculation is more clearly
defined in the male. '
87
88
BUCKETT
/. Res. Lepid.
This well marked speeies might be confused with P. liquida
Grote, or P. meodana (Smith), both of which possess a broader
subterminal space on the primaries than does lilacma. Also, the
subterminal line on the primaries of this species is not jagged
and irregular, but smooth, lacking the outward “W” mark so
prominently exhibited in liquida and meodana.
Another closely related species is P. rugosa ( Morrison ) , which
possesses an ash-grey pubescence covering the head, thorax, and
abdomen. A dark brown pubescence of the same areas and
more contrast in the markings readily separate lilacina from
rugosa. Greatest length of forewing in the male is 15 mm; of
the female, 16 mm.
Polia liquida (Grote)
Polia liquida Grote occurs in the northern coast ranges of the
western United States, extending northward into British Golum-
bia and eastward into Alberta in Ganada; in the United States,
it extends eastward into Idaho and Montana. Hampson (1905)
and Draudt (in Seitz, 1923) also list P. liquida in Colorado.
A male and a female were collected four miles east of Point
Arena, Mendocino County, 5 July 1958 (W. R. Bauer and J. S.
Buckett). Another pair of specimens was collected in the same
locality on 10 April 1959, by the same collectors.
Within this group, P. liquida may be confused with P. meo-
dana (Smith). Smith states, “The species (meodana) has been
confused with liquida Grt., which is a much more contrastingly
mottled form occuring in Washington . . . and as figured by
Hampson, has narrower, more pointed primaries . . . The
characteristics Smith observed in his specimens remain constant
in both species.
Greatest expanse of the forewing in the male is 15 mm; of the
female 16-17 mm.
Polia nevadae (Grote)
In the past few years, specimens of the rare Polia nevadae
(Grote) have been collected in fair abundance. Previous to
1960, this species was little known and poorly represented, if
not entirely lacking in most large collections. Forbes (1954)
records the “race” canadensis Smith, which is a lighter more
obscurely marked form of nevadae, from “New Brunswiek to
Alattagami River, Ontario, to Manitoba and Alberta, south to
Maine and Franklin Gounty, New York.” 1 have not studied the
types these names represent, but as Forbes is a lumper by most
standards, and yet considered canadensis to be a separate race,
there is reason to question the conspecificity of the forms.
7(2) : 87-94, 1968
FOLIA AND EUXOA
89
P. nevadae is a mottled brown, the basal area being orange-
brown above basal dash, otherwise basal and transverse areas
darker than subterminal area; a prominent cream colored “W”
mark can be seen on subterminal line; secondaries fuscous.
P. nevadae is quite a distinct species, its closest relative being
P. atlantica (Grote). The California specimens of nevadae can
not be too easily confused with atlantica, as nevadae is larger
and darker. The Colorado specimens of nevadae are identical
with, and conspecific with specimens taken near the type local-
ity. Forbes (op. cit.) states of atlantica “a pale race in Mani-
toba to Utah, often mistaken for nevadae.” Therefore, in speci-
mens taken from the Great Plains region, there is reason for
due caution in assigning a name to the collected specimens
believed to be either nevadae or atlantica.
The author has examined specimens from California and
Colorado, the majority being from Johnsville, Plumas County,
California, collected by Mrs. Helena Pini. Greatest expanse of
forewing 17-19 mm. P. nevadae was originally described from
the Sierra Nevada, California, Henery Edwards No. 4582.
Polia pulverulenta (Smtih)
Until recently, Polia pulverulenta (Smith) has not been re-
corded west of the Rockies. In general, it is northeastern in dis-
tribution. Its previously known distribution was the New Eng-
land States and adjacent parts of Canada, westward to Vancou-
ver Island, British Columbia. Crumb (1965) lists Colorado and
Washington also.
This species is dark ash-grey brown, perhaps its most distin-
guishing characteristic being a contrastingly large yellow lunule
in the tornus region of the primaries. Other California speices
most likely to be confused with pulverulenta are: 1) P. quadrata
(Smith), which is the same size or slightly larger, dark brown
in color and lacking the prominent yellow lunule in the tornus
region of the primaries; and 2) P. assiniilis (Morrison), also
closely related to quadrata but larger. Hampson (1903) syno-
nymized pidverulenta under assimilis but work of later authors
proved them to be separate species.
Eour specimens of pulverulenta were collected at the Uni-
versity of California Sagehen Creek Project, four miles north
of Hobart Mills, Nevada County, 21 June through 5 July 1962
by M. E. Irwin. One male is in the collection of the University
of California, Davis. Greatest length of forewing in both sexes
is 14 mm-16 mm.
90
BUCKETT
J. Rp.s. Lepicl.
Polia lutra glaucopis Hampson
Apparently, Polia lutra glaucopis Hampson is seldom recorded
from California, or from other western states. In the past year,
two California collections and one Oregon collection were re-
recorded. Two males were collected at Fort Dick, Del Norte
County, 16 and 30 April 1963, by J. W. Anderson, and one
female was collected 5 miles northwest of Corvallis, Oregon,
30 June 1962, by A. N. McFarland.
This subspecies was first discovered inhabiting Vancouver
Island, British Columbia, 2 females composing the type series.
P. lutra glaucopis was originally described as a subspecies of
P. luhens (Grote), but work of later authors prove lutra (Gue-
nee) and luhens to be conspecific, lutra being the older of the
two names.
This Polia is rather distinct and can be recognized by the
contrastingly light brown inner marginal half of the basal area
accompanied by its whitish tornus area and lilac subterminal
ar^a. The greatest expanse of the forewing varies slightly in
different specimens, the male being 18 mm, the female 20 mm.
The female of lutra lutra is slightly larger, with a forewing ex-
panse of up to 22 mm.
Barnes and Benjamin ( 1927 ) found . . no good character
to sort glaucopis, which is only western luhens, from typical
luhens. Western specimens are often darker in color than some
esatern specimens, but the character does not hold for any
series.” The fact that glaucopis is darker in coloration, accom-
panied with its more obscure markings and its smaller size,
will help to superficially distinguish it from the nominate sub-
speeies. McDunnough (1938) considers glaucopis as a sub-
species of Ultra, but future work will probably prove our western
subspeeies to be nothing more than a slight variant.
Euxoa extranea (Smith)
Heretofore, Euxoa extranea (Smith) has not been recorded
in California. With better collecting methods and increasingly
larger samples, extranea is now collected in large numbers,
enabling better evaluation of its specific variation.
California specimens of extranea differ slightly from typical
material in that the transverse anterior line of the primaries is
inwardly shaded and both the transverse posterior line and the
subterminal line are outwardly shaded with bands of cream-
colored scales. These shadings give the appearance of a lighter
ground color than that found in more northern material. This
differentiation might be confusing if only a limited number of
7(2) : 87-94, W68
POLIA AND EUXOA
91
specimens are available. Both sexes have dusky brown hind
wings rather than “yellow fuscous”, as stated in the original
description. Originally, extranea was described from a single
female collected in Montana, and perhaps with further study,
the California specimens will prove to be of subspecific merit.
In specimens of both sexes of extranea determined by Mc-
Dunnough (April, 1951, McD. No. 1144 Bauer-Buckett Collec-
tion), the ordinary crosslines of the primaries are black with no
shadings of cream-colored scales. These specimens were col-
lected at Mount Hood, Oregon, 17 August 1939, 6,000 foot ele-
vation (E. C. Johnston). In a series of over thirty specimens
from eight miles southwest of Johnsville, Plumas County, 11
August 1961 (W. R. Bauer and J. S. Bucket!), the maculation
of the primaries is quite consistently that of the lighter form.
Another specimen proving to be conspecific with extranea is a
female collected at Leavitt Creek, Mono County, 8,000 foot
elevation, 10 August 1960 (A. S. Menke). In this specimen, the
ordinary lines are as in typical extranea. Another specimen
from Hornbrook, Siskiyou County, 6 August 1961, is also typical
extranea.
One female is deposited in the collection of the University of
California, Davis. Greatest length of forewing in both sexes is
18 mm.
Euxoa vertesta (Smith)
Euxoa vertesta (Smith), a pale luteous species, was origin-
ally described from Stockton, Utah. E. vertesta is on the wing in
September and October over most of its range; however, speci-
mens have been collected from California in October only.
The available literature cites Utah as the only state in which
this species occurs. The author has before him twenty-six speci-
mens of both sexes from California, Nevada, and Utah. The
Calfornia series consists of six males and four females from
Olancha, Inyo County, 11 October 1962 (R. P. Allen). The
Nevada series consists of one male and two females from Pali-
sade, Eureka County, 4 September 1962 (T. R. Haig). The
Utah series consists of two males, one from Dividend, 6-17
September by the same collector.
E. vertesta may be confused with citricolor (Grote); however,
vertesta is characterized by its pale luteous coloration and weak-
ly defined reniform on the dorsal surface of the primaries. The
primaries of citricolor are light lemon yellow and the reniform
is more strongly defined. Both species possess white secondaries
and a white abdomen. Both species are found over much of
92
BUCKETT
/. Res. Lepid.
the same range . . . vertesta from Utah westward into California
and citricolor from Colorado and Arizona westward into Cali-
fornia and northward into Washington.
Greatest length of the forewing in both sexes is 15 mm. in
veriest a.
Euxoa edictalis (Smith)
Eiixoa edictalis (Smith) is typically of the Rocky Mountain
region in the United States, and extends westward through
Canada to British Columbia (Kaslo). Apparently, this species
has not been previously recorded from California, thus making
this large series before me from Alono County the first pub-
lished record.
E. edictalis occurs in the White Mountains, Alono County,
and can be collected quite abundantly in June at higher eleva-
tions. In 1962, and again in 1963, Mr. Paul Mannis and Mr.
David Mathais of the White Mountain Research Station, Mono
County, (elevation 10,150 feet), have ardently collected many
species of which edictalis was one.
Through the cooperation of Dr. David F. Hardwick, Canadian
National Collection, Ottawa, the author received specimens of
edictalis from both Colorado and from Kaslo, British Columbia.
From these specimens, close examination was made as well as
genitalic mounts of both sexes. This study proved the White
Mountain specimens to be very close to edictalis but there are
minor differences throughout.
E. edictalis is quite characteristic and can be confused with
no described species thus far. It is characterized by its deep
olive, grey-brown ground color of the primaries, the normal
markings being deep brown or black; the antennae of the male
are bi-pectinate; the thorax is very robust due to the great
amount of vestiture.
The greatest length of the forewing in the male is 16 mm; in
the female, it is 17 mm.
REFERENCES
CRUMB, S. E., 1956. The larvae of the Phalaenidae. 356 pp. Washing-
ton, D. C.
FORBES, W. T. M., 1954. Lepidoptera of New York and neighboring
states. Cornell Univ, Agric. Experiment Sta., Mem. 329. 433 pp.
GROTE, A. R., 1873. Descriptions of Noctiiidae primarliy from Cali-
fornia. Bull Buffalo Soc. Nat. Sci. 1:129-155.
HAMPSON, G. F., 1903. Catalogue of the Noctuidae in the collection
of the British Museum, vol. 4, London, 689 -}- xx pp.
, 1905. Catalog of the Noctuidae in the collection of the Brit-
ish Museum, vol. 5, London, 634 + ^vi pp.
7(2) : <S7-94,
FOLIA AND EUXOA
93
PLATE 1
Toi3 row, left, male, Polia lilacina (Harvey), Bishop, Inyo County, California, 3
August 1960 (R. P. Allen); top row, right, male, P. lilacina illahefacta (Morrison),
Fort Dick, Del Norte County, California, 25 June 1963 (J. W. Anderson); second row,
left, male, P. Jicfuicla (Grote), Fort Dick, Del Norte County, California, 21 May 1963
(J. W. A.); second row, right, male, P. incodana (Smith), Convict Creek, Mono County,
Califomia, 30 June 1963 (M. G. Timzi); third row, left, female P. riigosa (Morrison).
Ashland, Maine, 10 July 1945; third row, right, male, P. nevadae (Grote), Johnsville,
Plumas County, California, 7 June 1963 (H. J. Pini ) ; bottom row, left, female, P.
pulvcridcnta (Smith), Sagehen Creek, near Hobart Mills, Nevada County, California,
21 June 1962 (M. E. Irwin); bottom row, right, female, P. assimilis (Morrison), Lake
Katherine, Oneida County, Wisconsin (H. M. Bower).
HOLLAND, W. J., 1903. The Moth Book. Doubleday, Page and Co.,
New York, .xxiv + 479 pp., inehiding 48 plates.
McDUNNOUGH, J. H., 1938. Check list of the Lepidoptera of Canada
and the United States of America. Mem. Southern California Acad.
Sci., 1 (1): 1-272.
94
BUCKETT
J. Res. Lepiil.
PLATE 2
Top row, left, male. Folia lutra glacopis Hampson, Fort Dick, Del Norte County,
California, 16 May 1963 (J. W. A.); top row, right, female, P. quadrata (Smith),
Nelson Creek Road, 12 miles west Johnsville, Plumas County, California, 12 June
1961 (W. R. Bauer & J. S. Bucket! ); second row, left, female, Euxoa extranea (Smith),
Mount Hood, Oregon, 17 August 1939, 6,000 ft. elevation (E. C. Johnston); second
row, right, male, E. extranea (Smith)?, 8 miles southwest Johnsville, Plumas County,
California, 12 August 1961 (W. R. B. & J. S. B.); third row, left, female, E. citricolor
(Grate), 50 miles south Wells, Elko County, Nevada, 10 Sept. 1959 (T. R. Haig);
third row, right, female, E. vertesta (Smith), Olancha, Inyo County, California, 11
October 1962 (R. P. A.); fourth row, left, male, E. edictalis (Smith), Crooked Creek,
White Mountains, Mono County, California, 10,150 ft. elevation 26 June 1962 (J. S. B.
& G. M. Trenam); fourth row, right, female, E. edictalis (Smith), same data as pre-
ceding.
SEITZ, A. 1923. The Macrolepidoptera of the World, vol. 7. Stuttgart,
412 pp., 96 plate.s.
Journal of Research on the Lepidoptera
7(2) : 95-98, 1968
1160 W . Orange Grave Ave., Arcadia, California, U.S.A. 91006
© Coptjrighi 1968
VARIATION IN COLOR AND MACULATION IN A
POPULATION OF NEMORIA PULCHERRIMA
FROM THE SIERRA NEVADA OF CALIFORNIA
(LEPIDOPTERA: GEOMETRIDAE)
JOHN S. BUCKETT^ and TERRY A. SEARS^
^Systematic Entomologist,
California Department of Agriculture
Sacramento, Calif.
and
"Auhurn, California
It is our present intention to demonstrate the extreme vari-
ation exhibited by a population of the geometrid, Nemoria pul-
cherrima (Barnes and McDunnough), one of the greens from
Auburn, California. The specimens concerned were all collected
in the spring of 1967 on the rim of the American River Canyon.
The specimens were attracted to incandescent white light in
front of a white reflective surface from an area of mixed vege-
tation characterstic of upper Sonoran-transition zone localities.
The trees present were scattered individuals of Blue Oak ( Quer-
cus douglasii). Black Oak (Quercus kelloggii), Interior Live
Oak {Quercus wislizenii). Digger Pine (Pinus sabiniana), and
Ponderosa Pine {Pinus ponderosa). Undergrowth included
varied herbaceous vegetation and shrubs, the latter consisting
chiefly of Buckeye {Aesculus calif ornica) , Toyon {Heteromeles
arbutifolia) , and Manzanita {Arctostaphylos spp.).
The evenings on which the largest series were collected,
February 10 and 17, were fairly warm for that season and were
quite dark being one day after the new moon and the day of
the first quarter, respectively. Most moths came to the light
between the hours of 8:30 and 10:30 P.M., although some speci-
mens were captured as late as 11:30 or 12 P.M. None of the
95
96
BUCKETT AND SEARS
J. Res. Lepid.
specimens were reared; but it seems probable that the host
plant is oak as it was found to be in southern California by
Comstock and Dammers ( 1937 ) . The same host plant is cited
for piilcherrima by Comstock (1960). Whether or not he was
referring to his rearing of “naidaria” or whether he conducted
additional rearing experiments is unknown at this time.
Both the larva and pupa are described by Comstock and
Dammers (1937) under the specific name Nemoria naidaria
Swett. However, naidaria was synonomized under pulcherrima
by McDunnough (1938), and the condition remains the same
today. It might be interesting to note in McDunnough’s 1938
list that he cited “McD.” as author of pulcherrima. As the orig-
inal description was by Barnes and McDunnough, we have
adhered to the original citation, assuming the ''McD.” to be
an accident of some kind.
As can be seen by the type male in the Contributions ... by
Barnes and McDunnough (1916 vol. 3, pi. 2, fig. 10), this speci-
men most closely resembles our specimen which is the left one,
second row from the bottom on the colored plate. In recent
years a reddish form has been noticed in certain populations of
pulcherrima, and in a considerable percentage also. Of the
specimens examined from the 1967 collection at Auburn, one
third of the population was of the red form. Of the red speci-
mens only 10 per cent lacked the dark transverse lines of both
the primaries and secondaries; however, the black discal dots
seen in the upper left specimen (colored plate) were found to
be prominent. In the green specimens of the population, 60 per
cent of the specimens possessed prominent black transverse
lines, and 24 per cent possessed faint black transverse lines,
making up a total of 85 per cent of the green phenotypic portion
of the population. The discal dots were always found to a great-
er or lesser degree of prominence. In the coast range populations
of central California, one may encounter specimens in which
the discal dots may be entirely lacking, and seldom does one
collect specimens possessing the transverse lines.
It was suspicioned that the coast range populations and the
Sierran populations might represent two distinct entities speci-
fically, but there appears to be inadequate evidence at this
time to warrant the erection of a new name. Future revisionary
work and additional biological studies may reveal intricate char-
acters by which one may be able to distinguish the Sierran and
coastal populations at the specific level.
r
NEMORIA POPULATION
97
7(2) : 95-9H, 196H
98
BUCKETT AND SEARS
/. Res. Lepid.
REFERENCES
BARNES, WM., and J. H. McDUNNOUGH, 1916. Contributions to Rie
Natural History of the Lepidoptera of North America, vol. 3, pp.
1 - 296 + 33 pis. The Review Press, Decatur, 111.
COMSTOCK, J. A., and C. M. DAMMERS, 1937. Notes on the early
stages of three California moths. Bull. So. Calif. Acad. Sci. 36:68-78.
COMSTOCK, J. A., 1960. Inherent and applied camouflage in tlie sub-
family Geometrinae (Lepidoptera), including three new life history
studies. Trans. San Diego Soc. Natur. Hist. 12(26) :421-440, figs. 1-9.
McDUNNOUGH, J. H., 1938. Check list of the Lepidoptera of Canada
and the United States of America. Mem. So. Calif. Acad. Sci., No. 1,
pp. 1-272.
Journal of Research on the Lepicloptera
7(2) : 99-104, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
A NEW SUBSPECIES OF CALLOPHRYS
(CALLOPHRYS) DUMETORUM FROM
WASHINGTON AND OREGON
( Lycaenidae )
GLENN ALAN GORELICK
Vniversitij of California, Berkeley 94720
CALLOPHRYS ( CaLLOPHRYS DUMETORUM BdV., alODg witll
other members of this subgenus, is characterized by the green
color and the presence (or absence) and arrangement of white
maculations on the ventral surfaces of the wings. More than 500
specimens examined by the author indicate that this species
ranges from British Columbia to Baja California Norte. The
southernmost extension of this range, Los Angeles and San Diego
counties in California, and Baja California is supposedly in-
habited by C. dumetorum perplexa Barnes & Benjamin. Current
work on this subgenus by the author (in press under another
title), however, suggests that this subspecies is consubspecific
with typical C. dumetorum which occurs as far north as southern
Oregon. Unfortunately, problems exist concerning the identifica-
tion and classification of members composing the subgenus
Callophrtjs in that scale color on the ventral surfaces varies
from one population to another within a single subspecies. Life
histories of several populations of C. dumetorum were examined
to determine whether those populations in the Pacific Northwest
discussed by Clench (1963) are biologically distinct or in fact
variant populations of the typical form.
The biology of one population of typical Callophrys dum-
etorum studied by the author at Antioch, Contra Costa County,
California in April, 1967 was observed on a common leguminous
shrub, Lotus scoparius. This plant, common throughout the
Coast Ranges of California, is a known foodplant of this species
in southern California and females are known to oviposit on
Eriogonus fasciculatum foliolosum in the deserts ( Coolidge,
1924).
99
100
(.ORELICK
J. Res. Lepid.
Fig. 1 — Immature stages of CallopJirys dumetorum (Bdv.) — mature
larvae, prepiipa and pupa (V-18-1967) from Lotus scoparius Ottley.
( Brannan Island State Park, Sacramento County, California ) . ( Photo
courtesy of P. E. Turner, Jr.).
Fig. 2 — Mature, fourth instar larvae of Callophrys dumetorum oregonensis
Corelick on Lotus crassifolius. (Benth.) Green. (1964) (Falls City-
Valsetz Road area, Polk County, Oregon). (Photo courtesy of D. V.
McCorkle).
7(2): 99-104, 196H CALLOPHRYS DUMETORUM
101
In Washington and Oregon, available hostplant species diflFer
considerably. Newcomer (1965) lists Eriogomim heracleoides ^ E.
compositum and E. elatuni as hostplants for C. chimetorum, while
at Satus Pass, Yakima County, Washington, Dave McCorkle (in
correspondence - May, 1967) indicated that C. chimetorum
oviposits on Lotus nevaclensis. According to McCorkle (1965),
the Polk County (Oregon) population feeds on Lotus crassi-
folius, a common legume of the northern Coast Ranges. In
addition, it is interesting to note that specimens examined from
Benton County (Oregon) resemble typical C. chimetorum found
in California whereas several Polk and Yamhill County (Oregon)
populations have a phenotype similar to that of the Satus Pass
population. Although all of the plant species mentioned above
occur in California, they have never been recorded as hostplants
for C. chimetorum there.
Flight periods as given by Newcomer (1964) for Yakima
County, Washington show that the peak flight is in May, at
least one month later than the Antioch (California) population
as observed by the present author, with many southern California
populations often seen as early as February.
A comparison of the larvae from the Antioch population of C.
clumetorum and a Polk County (Oregon) population indicate
that they differ considerably in color and substrate (see photos
1 and 2 ) . On the basis of these life history differences and several
adult morphological details distinguishable from over 300 ex-
amined specimens of typical C. chimetorum from all over Cali-
fornia, a name was given to those Washington and northern
Oregon populations in the northern Coast Ranges.
Callophrys dumetorum oregonensis Gorelick
ssp. nov.
Holotype MALE; Costa of forewing 11 mm. from base to apex;
outer margin of forewing to CU2 and slightly indented at A2
margin of hindwing with shallow crenations, very weakly be-
tween Cui and Cu2, more pronounced between CU2 and
A2 as in the typical form; white annuli of antennae 15 (with an
incomplete 16th) as seen laterally; palpi dark above, with in-
termittent white scaling below; faeial hairs erect as in typical
form; body dark grave above, pale below; legs with both gray
and white scales, appearing annulated along tarsi.
102
GORELICK
/. Res. Lepid.
Dorsal surface of forewing a uniform gray rather than gray
brown seen in typical form; veins concolorous; stigma light,
greatly contrasting with wing; fringes of forewings and hind-
wings gray basally, becoming white apically.
Ventral suface with light green or yellow green scales reach-
ing posteriorly to Ciu; costa of forewing brown; fringes as on
dorsal surface.
Hindwings with green scales present over entire surface;
macular band present as two white spots with no apparent inner
black scales as soon in many of the paratype specimens; first
macule present in cell Sc, the second in cell Cm; fringes as on
forewings.
Allotype female: Differing from holotype male as follows:
White antennal annuli 17; dorsal surface uniform golden
brown; macular band on forewings present as two extremely
faint sports in Ma and Ciu cells, totally absent on hindwings.
Of a total of 55 paratypes examined, nearly all were con-
siderably smaller than the nominate subspeeies; the eosta of the
forewings in the latter is at least 13 mm. in length. The males
are gray, with little or no trace of brown, whereas females are
golden brown, often with gray seales present along the margins
of both the fore wings and hindwings. The fringes are always
with mixed dark and light seales as seen in typical C. diimetorum.
The green on the undersides of both sexes varies from light
green to grass green, most appearing much paler than the Cali-
fornia populations. The maculations on the hindwing undersides,
also quite variable in this subspecies, are present as several
separated faint spots, three or four closely connected bars, or an
incomplete macular band similar to that seen in C. sheridani
and C. viridis specimens. Most specimens examined show the
invasion of scales posteriorly to the Cus vein of the forewing as
mentioned earlier. Earlier descriptions of Callophrys (s. str. )
species used the term “fuscous” to defiine the brown scale shade
present; this term has not been used here in order to render
a more concise description.
Type locality: Kusshi Creek, 2200', Yakima County, Wash-
ton.
7(2): 99-104, 196H
CALLOPHRYS DUMETORUM
103
Type materials: Thirty males and twenty-seven females
as follows:
WASHINGTON, Klickitat Co,: Satiis Pass, 3000' to summit, ^ , V-4-55
(D. L. Bauer) $, V-18-63, 2 V-24-63, 9, VI-18-63, 2
9 , V-26-64 (all E. J. Newcomer), ^ , 2 9 , VI-8-63 (D. V. McCorkle).
Yakima Co.: 3 mi. E. of Fort Simcoe, 3 ^ , V-8-64, ^ , 9 , V-11-64, 4 9 ,
V-19-64 (all E. J. Newcomer); Kusshi Creek, 2200', 3 c? » 2 9 , V-24-63,
5,9, V-13-64, 2 ^ , V-23-64, S , V-21-65, ^ , V-9-66 (all E. J. New-
comer), (5, V-20-62 ( R. E. Woodley). Mason Co.: Shelton, $, 9,
V-23-57, 3 S, 2 9, V-2-58 (all D. L. Bauer); Stimson Creek,
IV- 17-49, ^ , V-7-49 (E. C. Johnston). Chelan Co.: Sand Creek, 2 9 ,
V- 29-57 (D. L. Bauer). Okanogan Co.: Black Canyon, $ , V-4-47 (E. C.
Johnston). OREGON. Clackamas Co.: near Big Eddy, 960', 9 , V-19-58
(no collector). Polk Co.: 4 mi. W. of Falls City, 1500', ^ , V-30-64
(D. V. McCorkle); Falls City-Valsetz Road area, <$ , 3 9, V-26-67
(D. V. McCorkle). Wasco Co.: 2 mi. SW of Rowena, 525', VI-6-64
(E. & S. Perkins); 15 mi. SW of the Dalles, 2600', $ , VI-16-62 (E. & S.
Perkins); 7.5 mi. E. of Bear Springs, Hwy. 52 at 3000', 9, VI-10-56
(O. E. Sette). Yamhill Co.: Baker Creek Valley, 300', ^ , 9 , VI-8-30, 9 ,
VI- 27-30 (K. M. Fender). IDAHO. Adams Co.: near mouth of Wildhorse
River, Wildhorse, 9 , V-12-59 (S. G. Jewett, Jr.).
The type material examined has been distributed as follows:
Holotype male and allotype female in the collections of the
California Academy of Sciences, San Francisco; two male para-
types in the collection of the California Insect Survey, University
of California, Berkeley; one male paratype and one female para-
type in the collection of the Los Angeles County Museum; one
male paratype in the collection of the Nevada State Museum in
Carson City; one male paratype in the collections of the U. S.
National Museum, Washington, D. C.; the remaining paratypes
are currently being retained in the collections of S. J. Jewett, Jr.,
D. L. Bauer, and the author.
I wish to extend my sincere thanks to E. J. Newcomer, Yakima,
Washington, David V. McCorkle, Monmouth, Oregon, Stan
Jewett, Jr., Portland, Oregon, E. Dornfeld, Corvallis, Oregon,
and David L. Bauer, Bijou, California for the loan of their speci-
mens without which this study could not have been undertaken.
104
GORELICK
J. Res. Lepid.
LITERATURE CITED
CLENCH, H., 1963. Calloplmjs ( Lycaenidae ) from the Pacific Northwest.
Jour. Res. Lepid. 2(2) : 151-160.
COOLIDGE, K. R., 1924. Life history studies of some California
Rhopalocera ( Lepidoptera ) . “The life history of Calloplmjs dumetorum
Boisdiival”. Trans. Am. Ent. Soe. 50(4) : 329-335.
McCORKLE, D. V., 1965. Contributor in: News of the lepidopterists’
society “Annual summary”. No. 3, page 5.
NEWCOMER, E. J., 1964. Butterflies of Yakima County, Washington.
“Callophnjs dumetorum, No. 68”. Jour. Lepid. Soc. 18(4) : 22^
NEWCOMER, E. J., 1965. Contributor in: News of the lepidopterists’
society. “Annual summary”. No. 3, page 4.
Journal of Research on Lepidoptera
7(2) : 105-111, 1968
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
& Copyright 1968
NOTE ON DAMAGED SPECIMENS
JOHN M. KOLYER
55 Chimney Ridge Drive, Convent, New Jersey 07961, U.S.A.
It is interesting to note the extent to which butterflies may
be damaged and yet remain capable of normal flight. This note
presents measurements on a few specimens with severe wing
damage and discusses these with reference to some of the per-
tinent literature.
ATTACKS BY BIRDS
There is considerable debate in the literature regarding bird
attacks, the interest being in supporting or discounting Batesian
mimicry. For example, Wheeler (1935) concludes that attacks
on flying butterflies are very rare and that most insectivorous
birds are incapable of capturing uninjured butterflies in flight.
This is stated to lead to the conclusion that “'the current theory
of mimicry as applied to the upper wing colors of butterflies is
unsound”. However, a considerable number (262) eyewitness
accounts of bird attacks compiled by Collenette ( 1935 ) showed
that 17% of the butterflies were captured at rest and 83% in flight.
Of course, it is recognized that in-flight attacks are the more
conspicuous, so that the only valid conclusion is that in-flight
captures are not uncommon.
Collenette ( 1935 ) also notes that symmetrical damage, as in
specimens 1 and 4 in the figure, strongly indicates a bird attack,
probably while the insect was at rest rather than with wings
momentarily together in flight. Carpenter (1942) examined
14,000 specimens for beak marks on the wings and concluded
that the small percentage of beak-marked specimens evidently
attacked at rest ( symmetrical damage ) militates against the view
that butterflies usually are attacked at rest. Therefore, it fol-
lows that mimicry on the upper surface would be perceived by
birds. This conclusion is in disagreement with Wheeler’s belief
105
106
KOLYER
J. Res. Lepid.
Fig. 1
All specimens were taken near Morristown, New Jersey and were flying strongly
when captured. 1 — Papilio polyxencs asterius Stoll, ^ , taken Aug. 6, 1966. 2 —
Papilio troitiis Linnaeus, g , taken Aug. 6, 1966. 3 — Papilio glaucus Linnaeus, ^ ,
taken Aug. 30, 1967. 4 — Limenitis arthemis astijanax Fabricius, ^ , taken July 30,
1966. 5 — Papilio troilns Linnaeus, ^ , taken Aug. 12, 1966. 6 — Hemaris thysbc,
taken Aug. 7, 1966.
7(2) : 105-111, 1968
DAMAGED SPECIMENS
107
that attacks on flying butterflies are rare but does no more than
remove one objection to the theory of upper-surface mimicry.
Incidentally, a recent eriticism of the eommon mimiery theory,
e.g. the Monarch-Viceroy relationship, is given by Urquhart
(1960).
An interesting conclusion by Carpenter (1942) is that attacks
by birds upon butterflies are predominantly (about 55% of cases
studied) from behind and less often from in front (about 30%)
or from the side (about 15%). Specimen 2 in the figure shows
what seem to be beak marks on the hind wings, while specimens
3 and 5 show considerable tearing; according to Collentte ( 1935 )
the majority of butterflies after being captured by birds show
torn wings rather than clear beak marks. However, as Collen-
ette notes, unless the attack is seen, torn wings cannot be ascrib-
ed to bird attacks with any degree of confidence.
Thus, specimens 1-5 seem to illustrate two cases of bird at-
tacks from the rear while at rest (specimen 4, which is clipped
very cleanly, and specimen 1), one case of bird attack(s) from
behind in flight (specimen 2), and two other possible cases of
attacks in flight ( specimens 3 and 5 ) .
EFFECT OF DAMAGE ON FLIGHT
The wing areas for the specimens in the figure were determ-
ined by inking the outline of the wings on clear plastic sheet
(0.042 inch thick), cutting along the lines, and weighing the
tracings with an analytical balance. The areas for undamaged
fore- and hindwings were determined similarly by consulting
undamaged specimens. The extents of wing areas remaining then
were calculated and are given in Table 1. Since all of specimens
1-5 were flying vigorously and apparently going about their usual
activities, a considerable part of the wing area ( 32% for specimen
1) seems to be expendable, at least when removed largely from
the hindwings.
Static loads (weight of insect divided by wing area) have
been reeorded for various insects; examples (experimentally de-
termined and from the literature) are given in Table 2. The
experimental determinations are based on weights of freshly-
killed specimens.
Assuming the static load to be about 0.009 g./cm.“ for an un-
damaged Papilio polyxenes asterius Stoll female, damage has
raised the load to 0.013 g./cm.- for specimen 1, an increase of
about 47% (neglecting weight of wing membrane lost).
108
KOLYER
J. Res. Lepid.
Table 1
EXTENTS OF DAMAGE FOR SPECIMENS 1-5
F or ewing,
P. C. Area Retained
Hindwing,
P. C. Area Retained
Total Area,
P. C. Retained
Specimen No.
Left
Right
Both
Left
Right
Both
1
96
83
89
43
50
47
68
Z
99
100
100
Z8
83
56
78
3
94
87
91
64
41
53
7Z
4
100
100
100
54
59
57
79
5
100
100
100
55
95
75
88
C. = per cent
Table Z
STATIC LOADS
Item
Papilio glausus Linnaeus (male)
Papilio troilus Linnaeus (male)
Cercyonis pegala Fabricius
Colias eurytheme Boisduval (male)
Hemaris thysbe
Papilionids and pierids
Butterflies in general
Static Load, g./cm. ^
0. 0093
0, 0081
0. 0063
0. 011
0. 10
approx. 0, 0 1
0. 01 - 0. 015
Bombus (Bombidae)
Monoplanes, circa 19Z6
0, Z5
1. 3 - Z. 3
7. 5 - 40
Reference
Portier, 1949
Portier and de
Rorthays, 19^6
Aircraft, circa 1953
Chadwick, 1953
7(2) : 105-111, 1968
DAMAGED SPECIMENS
109
The following simple experiments give some idea of the ex-
tent of wing loss that can be tolerated and of the relative im-
portance of forewings vs. hindwings.
Hemaris thysbe. — Complete removal of the hindwings (37%
of total wing area) had no apparent effect on flight, but removal
of the apices of the forewings ( comprising about 32% of the fore-
wing area), as shown in the figure (specimen 6), resulted in
slanting flight, perhaps 30° from the horizontal, toward the floor.
Limenitis archippiis Cramer. — After removal of the apices of
the forewings to the extent of 53% of the forewing area (27% of
total wing area), a specimen still was capable of level flight for
10 feet. The wingbeats seemed faster, as has been noted for in-
sects when the wing area is reduced (Chadwick, 1953). When
the hindwings (49% of total area) were quite removed, another
specimen flew well but somewhat erratically. Then, removal of
the apices of the same specimen to the extent of 20% of the fore-
wing area caused even more erratic flight, but level flight for
10 feet was achieved. The forewings (51% of total wing area;
forewings are 50-56% of the total for the four species of speci-
mens 1-5, incidentally) were quite removed from another speci-
men. Complete inability to fly resulted, and the insect was un-
able to rise above an inch from the floor. Thus, level flight is
possible using 80% of the forewings when the hindwings are
missing, but no flight is possible using 100% of the hindwings
when the forewings are absent.
Colias philodice Latreille. — As with the Viceroy, complete
removeal of the hindwings (51% of total wing area) from a male
specimen caused flight to be rather erratic, but the specimen
could sustain flight for at least one minute and was able to fly
across a 25 foot room and readily direct itself to a small (about
1 ft.-) window.
Papilio glaucus Linnaeus. — To test the effect of unsymmet-
rical damage, even more extreme than for specimen 5 in the
figure, one hind wing (25% of total wing area) was removed
from an undamaged female. Flight was not noticeably impaired.
CONCLUSION
In the Lepidoptera, the hindwings are said to associate closely
with the forewings to yield a single aerodynamic unit (Chad-
wick, 1953). However, though the wing area is about equally
divided between fore- and hindwings for the butterfly species
studied above, the forewings are dominant so that a limited part
110
KOLYER
J. Res. Lepid.
of the forewing area seems expendable vs. a major part of the
hindwings. Fortunately, attacks by birds tend to come from
the rear. Also, unsymmetrical damage can be tolerated, and
in-flight bird attacks, perhaps very common, tend to damage
the wings on one side more than the other (e.g., specimens 2,
3, and 5 in the figure).
It seems possible that the large wings of some butterflies are
a rather neutral factor in regard to survival of bird attacks. That
is, butterflies may be more conspicuous to birds than are bees,
for example, but an increase in relative frequency of attacks
may be balanced by reduced relative frequency of success in
that birds tend to peck at the partly-expendable wings (espe-
cially the hindwings) and miss the body. It might even be
hypothesized that the hindwings of certain species, for example
Papilio troilus Linneaus, which are conspicuously marked and
tailed, are of survival value in causing birds to peck at the most-
expendable part of the insect. Also, in many species the margins
(expendable) of both fore- and hindwings often are decorated
conspicuously.
This idea, like Batesian mimicry, might be most difficult to
demonstrate convincingly. Urquhart (I960) notes that bright
white tags applied to the wings of Monarchs seemed to attract
the attention of birds. A possible (though perhaps not practical)
experiment would be to apply white tags to various parts of the
wings of a large number of individuals of a suitable species
of butterfly and release these in a roomy aviary along with
insectivorous birds. To support the above ideg, signifiicantly
more specimens with tags on the hindwings and/or margins of
the wings should survive ( remain in flying condition ) than those
marked with tags on the inner parts of the forewings. A likely
result, of course, is that the birds might not be capable of enough
accuracy to strike at the particular part of the wing surface with
the tag. This would give survival rates independent of tag po-
sition and tend to discount the survival value of conspicuous
markings on the more-expendable areas of the wings, at least
for the particular bird species involved.
7(2) : 105-111, 196H
DAMAGED SPECIMENS
111
LITERATURE CITED
CARPENTER, G. D. H., 1942. The relative frequency of beak marks on
butterflies of different edibility to birds. Proc. Zool. Soc. London,
lllA (3/4): 223-231.
CHADWICK, L. E., 1965 The Motion of the Wings, in Insect Physiology,
edited by K. D. Roeder. fohn Wiley and Sons, Inc., New York. (pp.
582,591,599)
COLLENETTE, C. L., 1935. Notes concerning attacks by British birds on
butterflies. Proc. Zool. Soc. London, 1935(2): 201-217.
PORTIER, P. & (MLLE.) DE RORTHAYS, 1926. Load supported by the
wings in Lepidoptera of different families. Compt. Rend .Acad. Sci.
(Paris), 183(23): 1126-1129.
PORTIER, P., 1949. La Biologie des Lepidopteres. Paul Lechevalier,
Paris.
URQUHART, F. A., 1960. The Monarch Butterfly. University of Toronto
Press, Canada.
WHEELER, L. R., 1935. Do birds attack butterflies? Science Progress,
30(118): 272-277.
7(2) : 112, 1968
Journal of Research on the Lepidopfera
1160 AV. Orange Grove Ave., Arcadia, California, U.S.A. 91006
(P Copyright 1968
THE GENERIC, SPECIEIC AND LOWER
CATEGORY NAMES OE THE NEARCTIC
BUTTERELIES
PART 7 — The Genus Dryaduhi
PADDY McHENRY
1032 E. Santa Anita, Burbank, California
The single species occupying this genus has been shifted from
one genus to another since its original description by Linnaeus.
In addition to the genus Dn/adula Miehener, it has been included
by various authors in Papilio Linnaeus, Cethoski Fabrieius, Drijas
Hiibner, Colaenis Hiibner and Afjraidis Rosduval & LeConte.
The spelling of the name phaetusa has been unsatisfactorily;
Linnaeus, himself, spelled it first as phaetusa and later as
phaerusa. As phaetusa has definite priority it should be ac-
cepted as the proper spelling. The genera, including this one,
comprising the Nearctic subfamily Heliconiinae have an un-
fortunate history of spelling errors for the speeific names.
LIST OF THE GENERIC NAMES USED OR AVAILABLE FOR DRYADULA
DRYADULA Miehener.
Type, phaetusa (Linnaeus).
DRYADULA MICHENER 9 Oct. I94Z. Amer. Mus. Nov. (1197): 1,
no, 6; p. 4; figs. 5 and 10.
Type. P[apilio]. N[ymphalis]. [Phaleratus] phaetusa
Linnaeus. 1758. Syst. Nat. (10th. ed. ). 1: 478, no. 1Z3.
Type Selection. Miehener. 9 Oct. 194Z. Amer. Mus. Nov.
(1197): 4. He said: "Genotype, - Papilio phaetusa
Linnaeus, 17 58. "
LIST OF SPECIES AND LOWER CATEGORY NAMES USED OR AVAILABLE
FOR DRYADULA
1. DRYADULA PHAETUSA (LINNAEUS)
phaetusa (Linnaeus).
1. DRYADULA PHAETUSA (LINNAEUS).
phaetusa, P[apilio], N[ymphalis]. [Phaleratus] Linnaeus.
1758. ^st. Nat. p 0th. ed. ). 1: 478, no. 1Z3.
"Habitat in Indiis". No sex, series nor date
data given. The name was given as phaerusa by
Linnaeus. 1764. Mus. Lud. Ulr. ([!]): Z93, no.
111. The name was also given as phaerusa by
Linnaeus. 1767. Syst. Nat. ( 1 Zth. ed. ). 1(Z);
780, no. 180. Doubleday. 1847. List Spec. Lepid.
Ins. Coll. Brit. Mus. (Z): 65, misspelled the name
as pherusa and included it in the genus Agraulis.
112
Journal of Research on the Lepidoptera
7(2) : 113-121, 1968
1160 Orange Grove Avc., Arcadia, California, U.S.A. 91006
@ Copyright 1968
FIELD STUDIES OE CATOCALA BEHAVIOR
RONALD R. KEIPER
Department of Biology, Pennsylvania State University, Mont Alto
Although the Catocala have always been popular with
collectors, little information concerning their behavior and
ecology is available. Some observations on the natural resting
habits of adults have been recorded (e.g. Bunker, 1874; French,
1880; Johnson, 1882; Rowley & Berry, 1909; Kettlewell, 1958;
and Sargent & Keiper, 1969), but most of these are anecdotal
and not of a quantitative nature. The final paper is of particular
interest since it includes not only field observations of resting
Catocala, but also some experimental data which suggests that
at least one species of Catocala {Catocala antinympha) , along
with several non-Catocala species, are capable of selecting back-
grounds which match the reflectance of their fore wings. Before
field data can be used to substantiate these experimental findings,
it seems necessary to conduct field observations on a truly quan-
titative basis . Thus, to determine that moths actually do select
the appropriate background in nature, it must be shown that
they are not randomly selecting backgrounds, but instead are
actively choosing the appropriate ones from among a large
number of possible choices. This study presents some prelim-
inary work along these lines.
METHOD
An area of mixed forest was selected in Hampshire County,
in central Massachusetts, and the tree composition of the area
was determined. The actual number of each tree species, and
its percentage of the total forest composition, is presented in
Table L
Each day from July 15 to September 10, 1967, and from July
19 to August 5, 1968, the tree trunks of each of the trees on
the experimental plot were searched systematically for resting
moths, from ground level to a height of about 20 feet. When a
moth was found, it was photographed and extensive notes were
taken concerning the moth and its resting place. These notes
included information as to the species of tree selected, the resting
113
114
KEIPER
/. Res. Lcpid.
Table I. The tree species found on the study plot-their actual and
relative abundance.
Tree Species
Actual Number
on Plot
Percentage of the
total number of trees
on the plot
Black Birch
Betula lenta
48
33. 80
Red Oak
Quercus rubra
23
16. 20
Red Maple
Acer rubrum
21
14. 79
White Birch
Betula papyrifera
13
9. 15
White Oak
Quercus alba
10
7. 04
White Pine
Pinus strobus
9
6. 33
Hickories
Carya spp.
9
6. 33
Sugar Maple
Acer saccharum
7
4. 93
Hemlock
Tsuga canadensis
2
1. 41
Total Ntxmber of Trees
on study plot
142
7(2) : 113-121, 196H
CATOCALA BEHAVIOR
115
Fig. 1. Catocala rkUia “head-down” on black birch.
Fig. 2. Catocala concnmhcns “head-down” on white birch.
116
KEIPER
J. Res. Lepid.
height of the moth, and the resting attitude (“Head-up” or
“Head-down”). The moth was then captured in a glass jar, posi-
tive identification was made, and the moth was then released
back into the studv area.
RESULTS
A total of 70 moths of 14 species were found for which posi-
tive identification could be made and complete information
gathered. The total number of moths captured on each tree
species, and the percentage of this number to the total number
of moths captured, is shown in Table II. This data suggests
that there may be some selection occurring, for more moths
than expected are found on White Birch {Betiila paptjrifera) ,
and possibly Red Oak (Qiiercus rubra), while fewer moths than
expected are found on a number of tree species. This suggestion
thus requires closer examination of the distribution of each moth
species, and these results are included in Table III. Statistical
analysis, by Chi Square tests, show that only in the case of
Catocala relicta is there a significant difference between the
expected number of moths on a tree (White Birch) and the
observed number of moths. These results are particularly in-
teresting in that Catocala relicta is the only species of the 14
studied that is primarily white in color, and thus the only one
that would match the color of the bark of White Birch. The
other species, having darker forewings, would best match other
tree barks, and thus seem not to select particular backgrounds,
but only choose any relatively dark barked tree. There may,
however, be specific preferences or avoidances among species,
but the number of individuals within any one species is so far
too small for anv consistent trend to be determined.
RESTING ATTITUDE AND RESTING HEIGHT
Two other aspects of behavior have also emerged from this
study. First, certain species are consistent in their resting
attitude. Of the 14 species studied, only 3 species show a “Head-
up” resting attitude while the remainder rest “Head-down”.
Those species which rest “Head-up” are: Catocala relicta, C.
neogama, and C. iinijuga. Within any given species, this attitude
is consistent; in fact there were no exceptions among any of the
species. Thus all 7 of the C. relicta studied sat “Head-up”, while
7(2) : 113-121, 196H
CATOCALA BEHAVIOR
117
Fig. 4. Catocala relicta found at rest — “hcad-iip’' on white birch.
118
KEIPER
J. Res. Lepid.
Table II„ The distribution of the observed moths on the trees
of the study plot.
Tree Species
Black Birch (BB)
Red Oak (RO)
Red Maple (RM)
White Birch (WB)
White Oak (WO)
White Pine (WP)
Hickories (H)
Sugar Maple (SM)
Hemlock
Number of Moths
Captured
21
13
9
16
3
3
2
3
0
Percentage of
Total number of
Moths Captured
30. 00
18. 57
12. 86
22. 86
4. 28
4. 28
2. 86
4. 28
0. 00
Total number of moths
captured on study plot 70
Significant deviation from chance selection. . . Anaylsis by
Chi Square tests. Probability less than 0. 01.
7(2) : 113-12] , 1963
CATOCAf.A BEHAVIOR
119
all of the individuals of C. arnica (15), C. mclua (13), and C.
gracilis (12) sat “Head-down”. The importance of this resting
attitude is now being experimentally studied with respect to
possible functions associated with courtship or survival.
The second aspect of behavior noted was the consistency of
certain species to select certain resting heights, regardless of
the tree species rested upon. These results are shown in Table
IV. Species such as C. ilia seemed to show a definite preference
for resting high up on the trunk. This suggestion is further
substantiated by data from released moths. These moths had
been captured the previous night at “sugar”, kept overnight in
an experimental box (involved in other experiments), and re-
leased the next morning. Of 12 released C. ilia, 10 of the moths
selected resting places over 10 feet up on the trunks. On the
other hand, C. vidua seemed to prefer low resting places. Of
the 13 individuals observed in this study, all rested below 9
feet, with 7 of the number resting under 2 feet in height. Once
again, the importance of this behavior is not clearly understood,
but is being further investigated experimentally.
SUMMARY
A study was made of 3 aspects of Catocala behavior by ob-
serving resting moths in a woodlot in central Massachusetts
during the summers of 1967 and 1968. The first objective was
to attempt to determine whether moths selected then rested
upon backgrounds which tended to match their forewings. Al-
though the number of individuals in all cases was small, it ap-
peared that Catocala relicta, a moth with primarily white fore-
wings spotted with black to varying degrees, did select White
Birch for a resting place.
Secondly, the resting attitude was consistent within any one
species, but varied interspecifically. Three of the 14 species
studied rested “Head-up”, while the remainder sat “Head-down”.
Finally, there also seemed to be a preferred resting height
for a number of species. Some, such as Catocala ilia, generally
rested high up on the trunk, while others, such as Catocala
vidua, rested very low on the trunks.
These results then reveal that a high degree of consistency
exists in several aspects of Catocala behavior and suggest that
further study should be conducted to determine why and how
these unique behavioral responses occur.
120
KEIPER
/. Res. Lepid.
Table III. The distribution of the most commonly observed
species of Catocala on the trees of the study plot.
Catocala Species
BB
RO
BU
WB
WO
WP
H
SM
Total Number
of Moths
C. vidua
Z
3
3
4
0
1
0
0
13
C. relicta
0
0
0
6
0
0
1
0
7
C. concumbens
3
1
0
1
0
0
0
0
5
C« gracilis
5
0
1
4
0
1
0
1
IZ
C. ultronia
Z
3
1
0
1
0
0
0
7
C, arnica
6
5
1
0
1
1
1
0
15
. .
IZ
6
15
z
3
z
1
For abbreviations of tree species^ see previous table^ Table II.
Significant deviation from chance. . . Analysis by Chi Square
test^ Probability less than 0. 01.
LITERATURE CITED
BUNKER, R. 1874, Notes on collecting Catocalas, Canad. Entomol. 6: 25-
26.
FRENCH, G. H. 1880. Notes on Catocala hunting. Canad. Entomol. 12:
241-242.
JOHNSON, J. S. 1882. Catocalas taken in the vicinity of Frankford,
Pennsylvania. Canad. Entomol. 14: 59-60.
7(2) : 113-121, 1963
CATOCALA BEHAVIOR
121
Table IV. Resting Height of Eight Species of Catocala.
Data Refers only to Observed Resting Moths.
Resting Height
Catocala Species Low(0-3')
Medium (3-9’) High(Above 9')
C. vidua 8
5 0
C. ilia 0
0 3
C, relicta 2
3 2
C. unijuga 0
2 0
C.concumbens 1
3 1
C. gracilis 3
6 2
C. ultronia 0
3 4
C. arnica 3
9 3
Denotes those species which" rest '
'Head-up on the trunk.
(C. neogama also rest '’Head-up”,
but is not included in
the table since only 1 specimen was captured)
KETTLEWELL, H. B. D. 1958. The importance of the microenvironment
to evolutionary trends in the lepidoptera. The Entomol. 91: 214-224.
ROWLEY, R. R. and BERRY, L. 1909. Notes on the study of some Iowa
Catocalae. Entomol. News 20: 12-18.
SARGENT, T. D. and KEIPER, R. R. 1969. Behavioral adaptations of
cryptic moths. I. Preliminary studies on bark-like species. /. LepicL
Soc. ( In press).
Journal of Research on Lepkioptera
7(2) : 122
1968
HABITAT: SPECIFIC TYPE LOCALITY
Pleheius icariodes niissionensis H.
Slope on the west side of Twin Peaks, in the City of San Francisco,
California (Fig. 1). Approximately eighty percent of the area occupied by
this race was obliterated in the 1940’s by a housing development. This
race was once semi-continnoiis in distribution with P. icariodes pheres Bdv.,
the distribution of which was tied to that of the blue bush lupine in the
sand dunes to the west and north. The perennial prostrate lupine in the
foreground (Fig. 2) is the larval host plant. W. Hovanitz.
Jotirnal of Research on Lepidoptera
7(2) : 123-125, 1968
1160 W. Oranfic Grove Avc., Arcadia, California, U.S.A. 91006
@ Copyright 196H
LIFE HISTORY NOTES ON
SATYRIUM SYLVINUS DRYOPE EDWARDS
(LYCAENIDAE; THECLINAE)
THOMAS C. EMMEL and JOHN F. EMMEL
Department of Zoology, University of Florida, Gainesville 32601
and University of California Medical School, San Francisco
The hairstreak Satyrium sylvimis drijope Edwards is distribu-
ted through the coast ranges from the San Francisco Bay area
south to Los Angeles in California. S. s. drijope has been consid-
ered a species separate from sylvimis as late as Clench (1961);
however, the maculation and genitalia of the two entities are
essentially identical, with the only difference being that dryope
lacks the tail on the secondaries present in typical sylvinus (P. A.
Opler, in litt.; J. F. Emmel, in press). Its foodplant and life his-
tory have not been described ( Clench, 1961 ) .
The present paper presents a description of the first-instar
larva to make this information available for future comparative
studies of the larvae of the Theclinae, currently underway by
several authorities. The first-instar setal patterns seem to offer
the best differentiating characters among the hairstreak larvae
and doubtless will prove useful to ascertaining evolutionary re-
lationships when enough life histories are known.
GENERAL BIOLOGY AND FOODPLANT
Satyrium sylvinus dryope is single-brooded, with adults ap-
pearing in late May and June. The specific observations (during
1964-67 ) in this note are based on the dryope populations at the
Page Mill Road rock quarry on the Stanford University campus,
near Los Altos, Santa Clara County, and were made throughout
the flight season as well as at other times of the year.
123
124
EMMEL AND EMMEL
J. Res. Lepid.
\
/
/
Fig. 2
Fig.l. The first-instar larva of Satyrium sylmnus dryope Edwards, dorsal
view, with head and prothoracic shield to right and anal shield at left.
Fig. 2. Lateral view of the seventh body segment, showing setal arrange-
ment. Note the snpraspiracnlar round “gland” body.
7(2) : 123-125, 196H
LIFE HISTORY — SATYRIUM
125
The host plants are willows (Salix). Courtship of the adults
takes place around the willows bordering a wet seep. Eggs are
laid singly in willow bark crevices, especially at the junctions
of branches, in late May to early July. The species over-
winters in the egg stage. Larvae hatch the following March.
The first two instars feed by cutting a depression in the willow
leaf surface epidermis, rather than by cutting into the edge of
the leaf.
FIRST INSTAR LARVA
The body is flattened in the usual lycaenid shape. The spine
or setal arrangement is as shown in Figures I and 2.
The overall ground color is a uniform gray, with small dark-
brown elevated “dots” uniformly distributed over the entire
body surface. On the head-shield segment, a distinctly-shaped
yellowish-green area is outlined in brown. This head-shield area
lacks brown dots within it, but four spines protrude forward
from its margins.
On the anal-shield segment, there is a differently but distinct-
ly shaped yellowish-green area that is outlined in brown; this
area also lacks brown dots and it lacks spines.
The head is a dark brownish black in color and is kept hidden
while the larva is feeding.
The spines on all segments of the body and head are trans-
lucent, with a dark gray ring at the base of each.
Each body segment has a translucent, dark gray, raised
“gland”-like body or organ a short distance above the spiracle.
On the anal shield, near the anterior end, there are ten of these
dark gray “glands” arranged as shown in Figure I. Their func-
tion, if any, and homology with structures in the supraspiracular
position or elsewhere on other lycaenid larvae, remain unknown
(e.g., see Clench, 1962).
LITERATURE CITED
CLENCH, HARRY K., 1961. Tribe Theclini. In Ehrlich, P. R. and A. H.
Ehrlich, How to Know the Butterflies. Wm. C. Brown Co., Dubuque,
Iowa. 262 pp.
CLENCH, HARRY K., 1962. Panthiades m-album ( Lycaenidae) : Re-
marks on its early stages and on its occurrence in Pennsylvania. Journ.
Lepid. Soc., 15: 226-232.
EMMEL, J. F., in press. The genus Satyrium. In Howe, William H., edit..
The Butteiilies of North America. Doubleday & Co., New York.
Journal of Research on the Le))iflo]ifera
7(2) : 126
1968
HABITAT: GENERAL TYPE LOCALITY
Glaucopsijche lygdamus xerxes Bdv.
Plebejus icariodes pheres Bdv.
Sandy area near Lobos Creek, the Presidio, San Francisco, California,
the last known area for G. xerxes (Fig. 1). The xerxes host plant here is
Lotus sp. (Fig. 2 dried) and the pheres host plant is the blue bush lupine,
Lupinus chamisonis Esch. The last known collections here were made by
the author just prior to 1940. There have been no known collections since.
W. Hovanitz
journal of Research on ihe Le))i(loptera
7(2) : 127-130, 1968
1140 W. Orange Grove Ave., Arcadia^ California, U.S.A.
& Cnrn/righl 196S
THE GENERIC, SPECIFIC AND LOWER
CATEGORY NAMES OF THE NEARCTIC
BUTTERFLIES
PART 8 — The Genus Af'raulis
PADDY McHENRY
1032 E. Santa Anita, Bur])ank, California
In presenting this genus, I have followed, without approval
or disapproval, the name used for it by Dr. C. F. Dos Passos
(1964, p. 97) since he is the latest author to treat this group.
Many previous authors have considered that the type species of
Dione and A^miilis were congeneric and Dione has previously
enjoyed a popular usuage with the species vanillae although
Apraulis had still earlier enjoyed wide acceptance with it.
In addition to Dione and Agraiilis, vanillae has been associated
by various authors with the genera Papilio Linnaeus, Argynnis
Fabricius, and Dryas Hiibner.
Fabricius, and Dryas Hiibner.
The spellings for the specific names in this genus have been
relatively free of errors that seem to typify the other genera of the
subfamily Heliconiinae for the Nearctic area.
Agraulis 2/6
LIST OF GENERIC NAMES USED OR AVAILABLE FOR AGRAULIS
AGRAULIS Boisduval and LeConte.
Type, vanillae (Linnaeus).
dione HUbner ~
Type, juno (Cramer).
AGRAULIS BOISDUVAL and LECONTE. [1833] . Hist. Gfen. Icon. Lfepid.
Chen. I'Amer. Sept. 1(14): pi. 42; 1(16): 142-145. They
included only "Agraulis Vanillae".
Type. P[apilio]. N[ymphalis]. [Phaleratus] vanillae Linnaeus.
17^8. Syst. Nat. (10th. ed. ). 1: 482, no. 144.
Type Selection. As Agraulis vanillae was the only species
included in the genus by the authors, it became the
type.
127
128
McHENRY
J. Res. Lepid.
DIONE HUBNER. [1819]^. Verz. Bekann. Schmett. (2): 31, no. 4
He included only; "257. Dione Vanillae Linn. . . " and
"258. D. Juno Cram. . . " ^
Type. Pap[ilio]. Helicon[ius]. juno Cramer. [1780] .
Uitland. Kapellen. 3(24); 175. Described and figured
earlier without a generic name on page 38, as Fig. B-C
and on pi, 215, as figs. B-C in Pt^ 18, [ 1779]^.
Type Selection. Scudder. [8 Apr,] 1875 . Proc Amer.
Acad. Art Sci. 10; 157, no. 343. He said; "Juno may
be taken as the type. "
LIST OF SPECIES AND LOWER CATEGORY NAMES USED OR AVAILABLE
FOR AGRAULIS
1. AGRAULIS VANILLAE (LINNAEUS),
comstocki (Gunder).
fumosus (Gunder).
hewelettae (Gunder).
incarnata (Riley),
mar gineapertus (Gunder).
nigrior Michener.
passiflorae (Fabricius).
vanillae (Linnaeus ).
1. AGRAULIS VANILLAE (LINNAEUS).
comstocki, Dione vanillae Gunder. 5 Jan. 1925. Ent. News.
36(1); 5-6, no, 9; pi 1, fig. T. "Data; Holotype
d* ... Monrovia, Los Angeles County, California, July
19, 1924. "
fumosus, Dione vanillae Gunder. 4 May 1927. Ent, News.
38(5); 137, no. 9; pi. 2, fig. 9. "Data; Holotype?...
Los Angeles, Los Angeles County, California, Sept.
15, 1910..."
hewelettae, Dione vanillae incarnata Gunder, 6 Jan. 1930.
Bull. Brooklyn Ent. Soc. 24(5); 327-328; pi. 31, figs. 7.
"Data; Holotype $. . . Ontario, Riverside Co. , Calif.
Taken in the summer of 1927. . . "
7(2) : 127-130, 1968
GENUS AGRAULIS
129
incarnataj D[ioneJ . vanillae N. F. Riley„ Sept. 1926.
Entomologist. 59(760); 243-244. "Habitat; Southern
U. S. A. (Illinois, Texas, California and Florida, 9 dcf,
7 99, in B. M. ); Mexico (10 cTcf, 5 99, including types);
Honduras, 1 9; Guatemala, 1 d*, 2 99; Nicaragua, 4 d'd';
Costa Rica, 3 d'd*. The type cf and 9 are from near
Durango City, Mexico". No date data given.
margineapertus, Dione vanillae incarnata Gunder. 30 July 1928.
Canad. Ent. 60(7); 163; pi. A, fig. 3. "Data: Holotype
cT (fig. 3). . . Los Angeles. . .[California], Mar, 29, 1927.
In Author's coll. Two paratypes of similar immaculism
also in Author's collection.
nigrior, Agraulis vanillae Michener. 31 Dec. 1942.
Amer. Mus. Nov. (1215): 1-2 (in pt. ), in no. 2 (in key);
p. 7. "Holotype male: Upper Matecumbe Key, Florida,
February 19, 1932. . . ". "Allotype female; Indian River,
Florida, . . " "Paratypes, all from Florida. . ." Paratypes
from all months of the year except July, December and
January are noted. "The holotype, allotype and a
series of paratypes are in the collection of the
American Museum of Natural History. Additional
paratypes are in the collection of Mr. C. F. dos Passos
and Cornell University. "
passiflorae, P[apilio]. F[estivus]. Fabricius. [Between
“~TFAug. andTTDec. y IwT^Ent. Syst. 3(1): 60-61,
no. 189. "Habitat in Passiflora coerulea, laurifolia
Americes. . , " A new name for vanillae Linnaeus,
vanillae, P[apilio]. N[ymphalis]. [Phaleratus] Linnaeus.
1758. Sy7t. Nat. “(i 0th. ed.'). l7T82, nof 144.
"Habitat in Epidendro Vanilla Americes", No sex,
series nor date data given. Fazzini. 1934. Butt.
Moths Amer, : p, 8; fig. 21; gave name, in error, as
Dione vanille.
130
McHENRY
,/. Res. Lepid.
Agraulis 6/6
REFERENCES CITED
Dos Passos, C. F, 1964. A Synonymic List Of The Nearctic
Rhopalocera. Lepid. Soc. Memoir (1): i-vi^ 1-145,
FOOTNOTES
1. Dos Passos. 1959 [I960]. Jour. Lepid. Soc. 13(4)i Z12. Gave
additional date data for the Hist, Gfen. Icon.. Lfepid. Chen.
I'Amer. Sept, by Boisduval and LeConte.
2. Hemming. 1958. Official List Works Approv. Avail Zool.
Nomencl. (1): 4. Gave established dates for the Verz.
Bekann. Schmett.
3. Hemming 1958. Official List Works Approv, Avail. Zool.
Nomencl. (1): 9-10. Gave established dates for certain
works of Cramer and Fabricius,
4. A copy of the work among the separates at the Allan Hancock
Library (Univ. Sou. Calif. ) (ex library, Boston Soc. Nat.
Hist. ) has the following printed label on the front
wrapper: "Library of the Cambridge Entomological Club.
Received April 8, 1875, by gift from the author. "
5. The title page of Fabricius' Ent. Syst. , vol. 3, pt. 1 is
qualified by dates on pages [ii] and [488],
Journal of Research on the Lepidoptera
7(2) : 131-132, 1968
1140 W. Orange Grove Ave., Arcadia, California, U.S.A.
Coptfrighf 196H
DAYTIME VISION BY THE MOTH,
EXYRA RIDINGSI (RILEY)'
VERNON M. KIRK
Entomology Research Division, Agr. Res. Serv., USDA,
Brookings, South Dakota
Specimens of the Noctuid moth, Exyra ridingsi (Riley),
were observed by Jones (1904, 1907) resting within the leaves
of the pitcher plant, Sarracenia flava L., at Summerville, South
Carolina. In an area abounding in plants of this species, he
found that when a moth was dislodged from a leaf, it would
fly quickly to another leaf, alight outside near the rim, and
run in over the edge.
At 10:30 A.M. on 16 June, 1964, I found a specimen of E.
ridingsi within a leaf of S. flava growing in a grassy clearing
in a bog about 8 miles inland from Myrtle Beach, South Caro-
lina. When disturbed, the moth darted out of the leaf and
straight to the opening of a leaf of another pitcher plant 30
feet away. When again disturbed, the moth darted back to
the first plant, but into a different leaf. These 2 plants were
the only ones of this species within sight, and there was nothing
to block the view between them.
Further examination revealed 2 other specimens of the moth
in other leaves of the plants. When disturbed, these moths fol-
lowed a similar flight pattern in reaching the sanctuary of the
plant leaves, and in the 8 or 9 flights observed did not wander
more than 6 feet from a straight line between plants. Each flight
was completed in less than 3 to 4 seconds, indicating no hesi-
tance by the moths in choosing or locating their refuge.
1 Identified by Dr. E. L. Todd, IJSNM.
131
132
KIRK
J. Res. Lepid.
A careful search of the immediate vicinity revealed no other
specimens present, either on the ground or on vegetation. The
sky was clear; air movement during the period of flight obser-
vation was between 1 and 3 mph and at nearly right angles to
the flight path.
It appeared that direct vision was involved, although no
further attempt was made to test this possibility. If vision alone
were involved, it is remarkable that a moth is able to see and
identify, from a distance of 30 feet, a relatively low-growing
plant, in bright sunshine.
REFERENCES
JONES, F. M. 1904. Pitcher-plant Insects — I. Ent. News 15: 14-17.
1907. Pitcher-plant Insects — II. Ent. Neivs 18: 413:
NOTICES
BOOKS;
BUTTERFLIES. A concise guide in colour, Josef Moucha, ill. by
Vlastimil Choc. Paul Hamlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophieal
Library, N.Y.
WANTED:
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Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, California 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the Netv
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standing of this group of insects.
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THE JOURNAL OF RESEARCH
THE LEFUDORTERA'
IN THIS ISSUE
Studies on Nearctic Euchloe
Part 5. Distribution Paul Opler
Species in the Genera Folia and Eiixoa
John S. Buckett
Variation in Color and Maculation
in Nemoria pulcherruna
John S. Buckett and T. A. Sears
A New Subspecies of Callophrt/s dumetorum
G. A. Gorelick
Note on Damaged Specimens
John M. Kolyer
The Generic, Specific and Lower
Category Names of Nearctic Butterflies.
Part 7. The Genus Dryadula
Paddy McHenry
Field Studies of Cat oca la Behavior
Ronald R. Keiper
Habitat: General Type Locality,
Glaucopsijche hjgdamus xerxes
Plehejiis icariodes phercs W. Hovanitz
Life History of Satyriiim sylvinas dryope
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Habitat: Specific type locality,
Flebejus icariodes missionensis
The Generic, Specific and Lower Category
Names of Nearctic Butterflies.
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Daytime vision by the moth, Exyra
ridingsi Vernon M. Kirk
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Journal of Research on the Lepidoptera
7(3) : 133-148, 1968 (1970)
1160 W. Orange Grove Aoe., Arcadia, California, U.S.A. 91006
© Copyright 1968
FIELD WORK ON THE
POPULATION STRUCTURE
OF
OENEIS MELISSA SEMIDEA (SATYRIDAE)
FROM THE
PRESIDENTIAL RANGE, NEW HAMPSHIRE
GEORGE SGOTT ANTHONY
Dartmouth College,
Hanover, New Hampshire
INTRODUGTION
The purpose of this study was to assess the extent to which
various local populations of Oeneis melissa semidea are isolated
from one another. This subspecies of a characteristically North
American arctic and western alpine species is found throughout
the alpine areas of the Presidential Range in New Hampshire,
but as has been found with other butterfly species, it seems to
have certain localized areas of greatest abundance between
which individuals are seldom found. The presence or absence
of the butterfly in a given area of the range presumably depends
on the presence and quantity of available host food plant, in
this case alpine grasses and sedges, and various microclimatic
factors such as ground temperature, moisture, and depth of
snow cover during the winter.
The study was undertaken with the following two alternative
hypotheses as its basis:
1. The population of Oeneis melissa semidea is homogeneous
over the entire Presidential Range, that is, it is not broken down
into local breeding populations, and that a constant flow of
individuals and consequently genetic exchange occurs between
the various centers of abundance along the range. This situation
would tend to result in minimal or at least continuous variation
between samples of individuals drawn from selected areas of
the range.
133
134
G. S. ANTHONY
/. Res. Lepid.
bo
G
O
S
to
CO
=o
•2
"to
s
ments made in Roman numerals.
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
135
2. The population of Oeneis melissa semidea is not homoge-
neous, but rather it is broken down into discrete local breeding
populations with little or no exchange of individuals or genetic
information between them. This situation would tend to result
in greater variation between samples of individuals drawn from
selected areas of the range.
It was expected that the true situation for this butterfly lay
somewhere between the two hypotheses, depending on certain
ecological and environmental factors.
METHODS
The mark-release-recapture method was originally employed
to assess the movement of individuals between areas on the
range. Individuals were marked with dots of waterproof paint
(see Ehrlich, 1960), the position of which identified each indi-
vidual with a number. In this way, any individual recaptured
could be identified as to place of origin, and movements could
be detected. In addition, recaptures could be used to estimate
the size of the population as a whole, and for individual areas.
Between the dates of 27 June and 7 July, 51 marked individ-
uals were released in the Cowpasture, a rather extensive, rela-
tively flat area at mile 7 of the Mt. Washington auto road. No
subsequent recaptures of any of these individuals were made in
the Cowpasture or elsewhere. Since difficulty was being encoun-
tered in obtaining significant numbers of individuals for marking,
and because a number of factors related to the marking tech-
nique itself were becoming serious problems, the mark-release-
recapture attempts were ended, and pure sampling from the
population was begun.
From 8 July until 15 July samples of as large a number of
specimens as possible were taken from four selected areas of
the Presidentials. From north to south along the range these
areas were: Monticello Lawn on Mt. Jefferson (5300-5400 ft.),
the area surrounding the Gulf Tanks along the Mt. Washington
Cog Railway between the summits of Mt. Washington and Mt.
Clay (5700-5900 ft.), the Cowpasture at mile 7 of the Mt. Wash-
ington auto road (5700-5800 ft.) and Bigelow Lawn, directly
south of the cone of Mt. Washington (5400-5500 ft.). A total
of 115 individuals were taken by the author and another 30
were obtained from Donald Lennox of Jefferson, N. H., who
collected in the Cowpasture on 8 and 15 July. Of those collected
by the author, 13 females were kept alive and later released on
LOO — (n = 5) (n = l4) (n = 3l) (n=ll) (n = 58) (n = 23)
136
G. S. ANTHONY
4
h
4
J. Res. Lepid.
4
4
o
o
o
o
o
o
o
o
0)
00
1^
CO
m
ro
CM
d
d
d
d
d
d
d
d
Monticello Lawn Gulf Tanks Cowpasture Bigelow Lawn
Fig. 2. Frequency of spotting for samples of Oeneis melissa semidea from
areas of the Presidential Range sampled during the summer of_ 1969. Ob-
served frequency shown by horizontal dashes; 95% confidence limits shown
by vertical lines.
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
137
the summit of Mt. Mooselauke at the western edge of the White
Mountains. A grant total of 132 individuals were therefore
available for study following the initial field work required for
the study. Field work ended 2 August.
Up to the present time five characters have been analyzed in
an effort to provide evidence of genetic isolation between breed-
ing populations of the butterfly. Males and females have in all
cases been treated as separate populations in the analyses be-
cause of the lack of any evidence to treat them as the same, and
also because the mean values for each character analyzed differ
between males and females from a given area, sometimes signifi-
cantly.
The first character analyzed was the frequency of occurrence
of a spot in cell R5 of the forewing of the butterfly. Determina-
tion of the occurrence of a spot was made by visual inspection.
In only a small number of cases was the use of a hand lens neces-
sary for determination. The occurrence of a spot was defined
as the appearance of a group of scales between veins R5 and Mi
in the submarginal area of the forewing which were of a darker
color (usually black) than the ground color of the wing and
which were distinguishable on both the dorsal and ventral sur-
faces of the wing. This definition eliminated certain color vari-
ation appearing most commonly on the forewings of females
such as a small light ochreous patch of scales against a darker
ground color but without a darker center which is characteristic
of eyespots.
The remaining characters were the linear distances between
various points on the forewing ( see figure 1 for diagram ) :
I — extreme base of wing to the end of vein R4 (a standard-
ized indication of the overall length of the wing)
II — base of Mi to the end of R4 measured from inside discal
cell
III — width of discal cell from base of M3 to base of R3
IV — end of 2nd-A to end of R4 (indication of the width of
the wing)
Measurements were made under a 10 X dissecting microscope
with a scale accurate to 0.05 mm. This scale was made by photo-
graphing a 30 cm. ruler accurate to 0.5. and photographically
reducing the image 10 times on printing paper.
RESULTS
Results of the frequency of spotting analysis show a number
of interesting trends. First, the observed frequencies of the
occurrence of a spot in all cases were higher in female samples
138
G. S. ANTHONY
/. Res. Lepid.
Table 1
Values of "t scores" of comparisons of frequencies of spotting
between populations of Oeneis melissa from the Presidential Range^ N. H,
males
Monticello Lawn
Gulf Tanks
Cowpasture
Bigelow Lawn
females
Monticello Lawn
0.585
0.042
0.024
Gulf Tanks
0.610
0.671
Cowpasture
0.187
Bigelow Lawn
Monticello Lawn Gulf Tanks Cowpasture Bigelow Lawn
0.058
0.474
0.904
0.760
1.37
2.25*
* Significant
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
139
than in male samples from a given area. In only one case, that
of the Cowpasture samples, was the difference between male
and female frequencies significant (P<0.1). It is entirely possible
that, had sample sizes been larger, a significant degree of differ-
ence would have been found between male and female spotting
frequencies from the remaining three areas. This is especially
true for the Monticello Lawn and the Gulf Tankes areas. Com-
parisons of male samples showed a significant difference between
Bigelow Lawn and Cowpasture (P<0.025), and a difference ap-
proaching significance between Bigelow Lawn and Gulf Tanks
(P<0.2). All other comparisons of spotting frequency, including
those between female samples, showed no significant differences
(table I; figure 2).
Results of the forewing measurement analyses show that fe-
males are larger than males on the average, a result which is
almost invariable with most butterfly species (table 2; figure 3).
All but the Monticello Lawn samples show that this difference
is highly significant for measurements I and II ( table 3 ) . Again,
small sample sizes from Monticello Lawn may account for this
discrepancy. Between samples of a given sex from the four
areas very few differences even approaching significance were
found (tables 4 and 5). Males, however, tended to show great-
er differences than did females. Once again, little reliance can
be placed on values obtained from the male Monticello Lawn
sample because of its extremely small size.
DISCUSSION
The results of this study have been inconclusive. However,
the data from the five character analyses combined with field
observations can be used to draw at least tentative conclusions
until further field work can be undertaken.
From the observations of the butterfly, the hypothesis that the
population is divided into discrete local breeding populations
is likely, especially between the northern and southern portions
of the Presidential Range. Between Mt. Jefferson and Mt. Clay,
Tor example, the ridge drops to below 5000 ft. and enters typical
scrub vegetation. It is unlikely that semidea would fly this low
unless it were blown from higher ground. The butterfly rarely
flies more than a foot or two above the surface of the ground, so an
individual which might accidentally wander downslope would
eventually enter completely foreign vegetation and would proba-
bly seek higher ground again. This tendency to seek higher
ground, more commonly termed hilltopping, has been recorded
140
G. S. ANTHONY
/. Res. Lepid.
Table 2
Mean values and 95% confidence limits of wing measurements made on
samples of Oeneis melissa semidea from the Presidential Range, New
Hampshire.
Monticello Lawn
I
II
III
IV
male (n=5)
23.9 ± 0.99
10.4 ± 0.37
3.0 ± 0.18
14.6 ± 0.28
female (n=9)
24.4 ± 0.82
10.9 ± 1.5
3.0 ± 1.34
14.5 ± 0.34
Gulf Tanks
male (n=30)
23.0 + 0.39
10.1 ± 0.02
2.9 ± 0.09
14.1 ± 0.29
female (n=7)
24.3 ± 0.83
11.1 ± 0.73
3.0 ± 0.25
14.6 ± 0.50
Cowpasture
male (n=34)
23.2 ± 1.04
9.8 ± 0.59
2.9 ± 0.19
13.9 ± 0.95
female (n=12)
24.4 ± 0.57
10.9 ± 0.31
3.0 ± 0.09
14.4 ± 0.31
Bigelow Lawn
male (n=20)
23.4 ± 0.42
10.4 ± 0.38
3.0 ± 0.09
14.2 ± 0.34
female (n=9)
24.9 ± 0.97
11.3 ± 0.46
3.1 ± 0.19
14.8 ± 0.59
Table 3
Exact percentages (divided by lOO) of "t-scores" of comparisons
between male and female samples of Oeneis melissa semidea from designated
areas of the Presidential Range, New Hampshire.
ML
GT
C
BL
I
0.608
0.00505**
0.00069**
0.0015**
II
0.120
0.0006**
0.000**
0.0062**
III
1.000
0.326
0.080
0.243
IV
0.684
0.096
0.062
0.055
*
significant (P < 0.05)
highly significant (P < 0.01)
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
141
for many species of butterflies (for review see Shields, 1967)
including Oeneis melissa ( Munroe, 1948 ( 1951 ) ; Anthony, 1969 ) .
In the far north and the Rocky Mountains, Oeneis melissa flies
only above or north of the scrub line. On Mt. Washington and
the surrounding peaks of the Presidential Range the only indi-
viduals which actively fly appreciable distances are males in
search of females. Since the females almost invariably remain
in areas where grasses and sedges are the dominant form of vege-
tation, males tend to congregate in these areas also. Thus, be-
tween Mt. Jefferson and Mt. Clay a partial barrier to movement
of individuals and genetic exchange exists.
On Mt. Washington itself one interesting relationship seems
evident between the three sampled areas. Looking at the t scores
for spotting frequency computed between the samples from each
of the three areas, the significance of the differences between the
samples appears to be almost directly related to their directions
from each other relative to the direction of the prevailing wind
from the west. Between the Gulf Tanks and the Cowpasture
there is more than a 25% chance that the populations have the
same frequency of spotting. This is reasonable since the Cow-
pasture is in the direct path of the prevailing wind from the
Gulf Tanks. An individual could be and frequently probably is
blown from the Gulf Tanks area into the Cowpasture in a matter
of minutes. On the other hand, between the Gulf Tanks area
and Bigelow Lawn there is only about a 15% chance that the
populations have the same frequency of spotting. Again the di-
rections of Bigelow Lawn from the Gulf Tanks relative to the
direction of the prevailing wind would account for the reduced
chances that the populations have the same frequency. Finally,
between the Cowpasture and Bigelow Lawn there is almost no
chance, less than 2%, that the populations have the same fre-
quency of spotting. This is because there is almost no chance
that an individual could be blown from one to the other, since
a line between them is practically perpendicular to the direction
of the prevailing winds. Individuals which are blown anywhere
are probably blown directly east or southeast, in the case of the
Cowpasture into the Great Gulf or Huntington Ravine, and in
the case of Bigelow Lawn into Tuckerman’s Ravine or the Gulf
of Slides. In addition, the Alpine Garden, which lies directly
between the Cowpasture and Bigelow Lawn, is surprisingly
devoid of the butterfly, even though in the past it has been
regarded as a prime collecting area for Oeneis melissa.
142
G. S. ANTHONY
/. Res. Lepid.
6
£
Fig. 3; Diagram of values of measurements made on the forewings of
Oeneis melissa seniidea from the Presidential Range, N. H. Roman num-
erals correspond to those of fig. 1. ML = Monticello Lawn, GT = Gulf
Tanks, G = Gowpasture, BL = Bigelow Lawn.
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
143
Table 4
Exact percentages (divided by 100) of "t-scores*’ of comparisons
of wing measurements made on samples of Oeneis mellssa semidea from the
Presidential Range j New Hampshire.
Males
ML
GT
C
BL
Measurement
ML
0.078
0.199
0.283
(1)
-- -
0.249
0.038*
1.000
(11)
0.062
0.301
1.000
(111)
—
0.159
0.126
0.251
(Iv)
GT
—
0.567
0.126
•—
0.030*
0.122
1.000
0.115
— -
0.323
0.652
C
0.543
—
0.002**
>—
0.050*
0.164
BL
^ significant
highly si ^lif leant
144
G. S. ANTHONY
/. Res. Lepid.
Table 5
Exact percentages (divided bj 100) of "t-scores" of comparisons
of wing measurements made on samples of Oeneis melissa semidea from the
Presidential Range^ New Hampshire,
o
Female s
ML
GT
C
BL
Measurement
ML
—
0.840
1.000
0.622
(1)
—
0.586
1.000
0.175
(11)
___
1.000
1.000
0.657
(111)
—
0.709
0.748
0.639
(IV)
GT
0.814
0.308
(V)
—
0.506
0.577
(VI)
—
1.000
0.537
(vil)
—
0.575
0.589
(VIII)
C
—
0.298
(IX)
0.100
(X)
—
0.268
(XI)
— -
0.172
(XII)
BL
—
(XIII)
—
(XIV)
(XV)
(XVI)
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
145
Two points mus be emphasized in defense of this attractive
relationship. First, the period of time during which this butter-
fly flies during the summer is short, beginning slightly before the
first of July with very small numbers of individuals, reaching a
peak in numbers by the end of the second week of July, and
ending by the last week in July or the first of August, depend-
ing on the weather conditions for a given season. This year
( 1969 ) , the flight period was probably effectively ended by the
22nd of July because of extremely bad weather which began on
that date. By the 2nd of August, when the harsh weather had
ended, no butterflies were seen anywhere. Since the flight period
of the butterfly is therefore so limited, the period during which
genetic exchange between local populations is possible is limited
as well Secondly, during this short flight period the number of
days during which adults fly is limited by the weather. The
butterfly tends to fly in appreciable numbers only on relatively
warm, sunny days, which are few and far between in this above
tree-line area of the White Mountains. The butterfly generally
will not fly in winds above 40 mph., in temperatures below 45
degrees F., or in fog or rain, unless it is disturbed. Since the
butterfly then only flies during relatively ''good” weather, and
since the wind rarely blows from directions other than the west
or northwest during such ‘“good” weather, the chance of move-
ment of individuals by wind action alone in any direction other
than from, west to east is slight.
A number of inconsistancies exist in the data and field obser-
vations of Oeneis melksa semidea. First of all, comparisons of
frequency of spotting made between areas are not paralleled by
the. comparisons of wing measurements. In fact, the lack of a
definite pattern to the comparisons of the wing measurements
casts doubt on the validity of these measurements as genetically
controlled characters. The fact that many species of Oeneis
possess a spot in exactly the same area of the wing that certain
individuals of semidea do would seem to indicate that spotting
is indeed genetically controlled and not subject to differences
in time of eclosure, nutritional factors, or other environmental
conditions. However, overall size of the butterfly, reflected in
the measurements, may indeed be influenced by the above fac-
tors. Breeding experiments should resolve this question if a
successful technique for raising the butterfly can be developed.
Another inconsistency is found in comparing male and female
spotting frequencies from a given area and between areas, and at
the same time recalling that the females of this species are more
146
G. S. ANTHONY
J. Res. Lepid.
New Hampshire.
Areas sampled for Oeneis melissa semidea indicated by large capital
letters, inclosed by dashed line. ML = Monticello Lawn, GT = Gulf Tanks,
C rz Cowpasture, BL zz Bigelow Lawn.
7(3) : 133-148, 1968 (1970)
POPULATION STRUCTURE
147
or less sedentary, whereas the males tend to wander. In this
case the question arises as to why the female spotting frequencies
are statisically constant and do not at least parallel the males.
The only explanation for this that so far appears to be tenable is
that the males, because they fly more often and for greater dis-
tances are therefore more exposed than the females and are
consequently subject to some unknown selective pressure more
than are the females. This selection pressure may be resulting
in more variation in the male populations than in the female.
An interesting point to mention is that there seems to be a slight
correlation between the presence of a spot on a given individual
and the distinctness of the dark median band on the ventral
surface of the hindwing. If indeed there is a correlation between
these two characters, then a basis can be laid for suspicion of
predatory pressure favoring spotted individuals through selec-
tion for a distinct band. It has been proposed that the white
band which crosses the wings of certain species of Limenitis
butterflies produces a form of disruptive coloration, breaking
up the outline of the wing and rendering it less easily seen by
a potential predator (Platt and Brower, 1968). It seems pos-
sible that adult semidea are under selective pressure from preda-
tors such as certain species of birds which frequent the alpine
areas of the Presidential, and that the distinctness of the median
band of the hindwing (which is exposed when males sun them-
selves on rocks ) and likewise the presence of a spot on the fore-
wing is being influenced by this pressure. It is also interesting
to note that another species of Oeneis, namely polyxenes, is
found in a very similar arctic relict environment on Mt. Katah-
din in Maine. All individuals of this population possess a spot in
exactly the same area of the forewing and all are rather invari-
ably distinctly banded. Oeneis melissa from the Presidential,
on the other hand, vary considerably in not only the presence
but the overall development of a spot as well as in the dis-
tinctness of the median band. Unfortunately, no objective
means could be devised for determining whether or not a band
is distinct, and hence the correlation between band and spot has
yet to be statistically shown.
SUMMARY AND CONCLUSIONS
Oeneis melissa semidea from the Presidential Range of New
Hampshire was studied in the field, and samples from four areas
of the range were taken in an effort to determine the popula-
148
G. S. ANTHONY
J. Res. Lepid.
tion structure of the butterfly. Statisical treatment of five char-
acters yielded no conclusive evidence for either total isolation
or lack of isolation between the populations inhabiting the four
areas, but field observaton combined with the statistics derived
from the frequency of the occurrence of a spot on the forewing
of the butterfly indicate that at least partial barriers probably
exist between the sampling areas. Movement of individuals be-
tween any of the areas was not seen while in the field. Move-
ment by action of the prevailing wind from the west is dis-
cussed and cited as probably the major contributor to the break-
down of any spatial or environmental barriers which do exist.
ACKNOWLEDGMENTS
I wish to thank Dr. Andrew Nelson and Dr. John Gilbert for
their guidance and assistance in planning and carrying out this
undertaking, the crew of the Mount Washington Observatory,
especially Guy Gosselin, the Chief Observer, for their hospital-
itl and company, and the Undergraduate Summer Research
Fellowship Committee of Dartmouth College for making this
summer’s work possible. This work was supported by a fellow-
ship derived from PHS grant number 5 TOl HE 5303-11.
LITERATURE CITED
ANTHONY, GEORGE SCOTT, 1969. Field notes and subspecific status
of Oeneis melissa (Satyridae) in northern Quebec. Jour. Lepid. Soc.
23: 103-104.
EHRLICH, PAUL R. and SUSAN E. DAVIDSON, 1960. Techniques for
capture-recapture studies of Lepidoptera populations. Jour. Lepid.
Soc. 14: 227-229.
MUNROE, EUGENE, 1948 (1951). Field notes on the butterflies of Knob
Lake, Northern Quebec. Lepid. News 5: 7-10.
PLATT, AUSTIN P., and LINCOLN P. BROWER, 1968. Mimetic versus
disruptive eoloration in intergrading populations of Limenitis arthemis
and astyanax butterflies. Evolution 22: 699-718. ( 1
SHIELDS, OAKLEY, 1967. Hilltopping. An eeological study of summit
congregation behavior of butterflies on a southern California hill.
Jour. Res. Lepid. 6(2): 71-178.
Journal of Research on the Lepidoptera
7(3) : 149-152, 1968 (1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
A HYBRID LIMENITIS FROM NEW YORK
ARTHUR M. SHAPIRO
and
JAMES D. BIGGS
Department of Entomology & Limnology^ Cornell University,
Ithaca, New York,
14850
An apparent hybrid between Limenitis arthemis Drury and
L, archippiis Cramer ( Nymphalidae) was taken by one of us
(AMS) on 6 August 1967 in Dryden Township, near Ithaea,
Tompkins County, New York. The speeimen (figs. 1, 2) is a
male in fresh eondition. It agrees in most respeets with the pub-
lished deseription of L. X arthechippus Seudder (1889), and
the upper surfaee is nearly identieal to one of the bred examples
figured by Field (1914).
There are at least four prior records of this hybrid from the
Northeast, plus one involvng the northwestern subspecies of
L. arthemis, rubrofasciata B. & McD. The type specimen was
first described by Edwards (1882) as L. arthemis, ab.C. The
same specimen later served as the type of Scudder’s arthechip-
pus. It was collected at Chateauguay, Que. (vie. Montreal) by
J. G. Jack in 1879. The other Eastern records are all from Field,
who collected somewhat darker specimens at Alstead, N. H.
in 1895 and 1896, and saw another in 1902 (Field, 1904). The
Western specimen was collected at Beulah, Manitoba, and
described as L. X rubrofasechippus by Grinder (1934). There
seem to be no recent records from the East.
The somewhat similar L. X rubidus Strecker, a putative hybrid
of L. archippus and L. astyanax Fabr., is known from Berks Co.,
Pa.; Jeannette, Westmoreland Co., Pa.; Wellesley, Mass.; “Shar-
born” (Mass.?), Brooklyn, N.Y.; and “eastern N. Y.” (probably
Catskills). A recent example collected at Louisville, Ky. was
described by Monroe (1953).
149
150
SHAPIRO & BIGGS
J. Res. Lepid.
Fig. 1. Limenitis from Tompkins Co., N. Y. Upper surfaces. Top Left:
L. arthemis $ , Cayuga Inlet Valley, 3.viii.67 (AMS). Top right: L.
archippus $ , Monkey Run, Dryden, 6.viii.67 (AMS). Bottom: L. X
arthechippus $ , Monkey Run, Dryden, 6.viii.67 (AMS).
151
7(3) : 149-152, 1968 (19W) HYBRID LIMENITIS
I
1
'I
Fig. 2. Lower surfaces of the specimens shown in Fig. 1.
152
SHAPIRO & BIGGS
J. Res. Lepid.
The sexes of the rubidus are not known, but all of the arthe-
chippus recorded are males. Field (1914) reported an experi-
mental cross of reared female archippiis X wild male arthemis.
He obtained poor egg hatchability ( 19/ 62 ) and an abnormal sex
ratio (8^ :0$ plus a dead pupa probably male; for 9:0 with
expected l:lx^=9.0, P< .005). The preponderance of males is
in accord with Haldane s Rule. Other broods of hybrid Limen-
itis, reared by Remington, also show this phenomenon (Rem-
ington, 1958).
The very different coloration of L. arthemis and L. archippus
would suggest the existence of strong behavioral barriers to
hybridization. Through most of their range the two species are
strongly, but not totally, isolated on an ecological basis, arthe-
mis being essentially a woodland insect while archippus occurs
principally in open country. In this connection it is of interest
that the Ithaca hybrid was taken in a disturbed, ecotonal area
in close proximity to typical habitats frequented by the parent
species. On the same stand of Teasel (Dipsaciis sylvestris
Huds.) with the hybrid were several normal archippus, one of
which is figured, while in the woods several hundred feet away,
fresh arthemis of the second brood were flying. The known
food plants of L. arthemis near Ithaca are Populus tremuloides
Michx. and P. deltoides Marsh. L. archippus has been reared
locally on P. deltoides and observed ovipositing on willows
(Salix). Of these, Salix spp. & P. tremuloides were present in
the vicinity of the collection site.
LITERATURE CITED
EDWARDS, W. H., 1882. Description of new species of butterflies
found in the United States. Papilio 2:45-49.
FIELD, W. L., 1904. Problems in the genus Basilarchia. Psyche 11:1-6.
1914. Hybrid butterflies of the genus Basilarchia. Psyche 21:
115-117.
GUNDER, J. D., 1934. A check list revision of the genus Basilarchia
Scudder. Canadian Entomologist 66:39-48.
MONROE, B. L., R., 1953. A hybrid Limenitis. Lepidopterists News
7:53.
REMINGTON, G. L., 1958. Genetics of Populations of Lepidoptera.
Proc. X Int. Cohgr. Ent. 2:787-806.
SGUDDER, S. H., 1889. Butterflies of the Eastern United States and
Canada. Vol. 1, p. 296. Author, Gambridge, Mass.
Journal of Research on the Lepidoptera
7(3) : 153-165, 1968 (1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
THE POPULATION BIOLOGY OF THE
NEOTROPICAL SATYRID BUTTERFLY,
EUPTYCHIA HERMES.
I. INTERPOPULATION MOVEMENT, GENERAL ECOLOGY,
AND POPULATION SIZES IN
LOWLAND COSTA RICA (DRY SEASON, 1966).
THOMAS C. EMMEL
Department of Zoology, University of Florida, Gainesville 32601 »
No THOROUGH STUDY OF LEPIDOPTERAN POPULATION STRUCTURE
in the Neotropics has been done, yet the butterflies reach their
greatest diversity in this biogeographic realm. The present in-
vestigation of Euptychia hermes Fabricius (Satyridae) is the
first to involve a neotropical satyrid butterfly, and because the
species ranges north to the Atlantic coast of the United States
the choice of this Euptychia will allow future comparisons of
the structure of both temperate and tropical populations of the
same species. Reported here are data obtained on population
size and intrapopulation and interpopulation movement in popu-
lations of E. hermes located in western and eastern Costa Rica,
in Central America.
MATERIAL AND METHODS
Euptychia hermes is a small satyrid, dull brown in ground
color, and with a wingspread of about 25 mm. The sexes are
similar in coloration and pattern. On the undersides of the
wings are several reddish-brown lines (submarginal and limbal
positions ) and a number of marginal ocelli. The ocelli are rather
indistinct on the forewings, but on the hindwings, six well-
marked ocelli are present (Figure 1). The degree of develop-
ment of these hindwing ocelli was used as an index to phenetic
variation (data to be reported later).
153
154
T. C. EMMEL
J. Res. Lepid.
Fig. 1, Euptychia hermesi ventral surface, showing ocellation pattern.
7(3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
155
A capture-mark-release-recapture program to determine popu-
lation size and extent of individual movement was carried out
in a series of populations in Guanacaste Province (four success-
ive days) and in one population in Limon Province (six suc-
cessive days), Magic-Marker ink pens were used to mark the
individual butterflies on the wings by a code system (Ehrlich
and Davidson, 1961). As this butterfly was found to be most
active in the early morning, all marking studies were done be-
tween 7:00 and 9:30 a.m. and repeated daily. Each area was
sampled once a day to avoid the complication of same-day
recaptures. All butterfles flying in a population could be cap-
tured in 30 to 40 minutes at most, and the earlier captures were
retained in extra nets until that time when all specimens were
marked (or recorded if a recapture) and then released again.
DESCRIPTION OF STUDY AREAS
1. Guanacaste Populations (Western Costa Rica):
A series of five populations were located along a thousand-
foot ( 300-f- meters ) section of the river road east of the experi-
mental station of the Costa Rica Ministry of Agriculture and
Livestock, adjacent to the Finca Taboga, located 13.5 km.
southeast of Cahas, Guanacaste Province, at an elevation of 11.5
meters (38 feet) above sea level. The study was carried out
February 13-16, 1966.
The general vegetational formation was dry tropical deciduous
forest, in a late stage of second-growth recovery along the road
where a series of grassy "islands” were surrounded by vine-
covered shrubs and dense undergrowth except on the road side,
and were isolated from each other by differing expanses of dry
habitat. The small meadows were bordered on the east by a
swamp, the source of moisture for the green grasses. Euptychia
hermes at this location was abundant in these grassy areas; in-
dividuals were occasionally encountered elsewhere in the dry
forest. The entire group of colonies in the study area was isolated
from other grassy regions along the road by at least 250 feet
(70 meters). Figure 2 shows the spatial orientation of the
population sites.
The grasses were one-third to one-half meter in height; the
surrounding vines and shrubbery were 1.5 to 2 meters and more
in height. The butterfles flew over these latter "barriers” on
occasion, but usually flew into them through slight "holes” in
the leafy wall, when pursued or when the sun rose higher in
the late morning. Second-growth plants found in the study area
156
T. C. EMMEL
J. Res. Lepid.
Fig. 2. The spatial orientation of the population sites studied along the
river road on the Finca Taboga, southeast of Cahas, Guanacaste Province,
Costa Rica.
Fig. 3. Map of the population areas studied on the Los Diamantes ex-
perimental station grounds, east of Guapiles, Limon Province, Costa Rica.
Fig. 4. Daily fluctuation in flight activity of Euptijchia hermes at the low-
land Costa Rican study sites: open meadow populations.
7(3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
157
were: Solanum (3-4 meters high, a border tree), two species of
Salvia (Labiatace), Oenothera, Cassia (a border tree), several
species of vines ( Convolvulaceae and Vitaceae), Philanthus
( Euphorbiaceae, a border plant), Tripleris (a border tree),
Panicum grass) and two unidentified grasses (one of which the
butterfly flew around and frequently landed on).
The daily weather here was sunny and hot with intermittent
clouds. During the hours of the capture-recapture studies, the
temperature ranged from about 75 to 86°F., a daily maximum
of 96 to 98° F. was usual. The relative humidity was around
50%. A strong gusty wind developed by 9:15 a.m. every day.
IL Los Diamantes Populations (Eastern Costa Rica):
The capture-recapture study was done March 4-9, 1966, on
a population inhabiting a lush grassy area surrounded by cleared
fields and cacao and rubber plantations (these areas with grass-
es also), located about a half kilometer northeast of the build-
ings at the Los Diamantes experimental station of the Costa
Rican Ministry of Agriculture and Livestock, 1 km. east of
Guapiles, Limon Province, at an elevation of 300 meters (984
feet). An adjacent population was sampled to study local vari-
ation patterns (see Figure 3 for map).
The grasses were lush on the east side of the road and up to
35 cm. in height; many Euptijchia were flying here. On the west
side of the road, the grass had been cut short and few butterflies
occurred there. In Area C, many Euptychia were landing on
crushed sugar cane stalks in the road and sipping the sap. The
peak of butterfly activity, as at Guanacaste, was between 7:00
and 9:30 a.m. The daily weather was warm, from about 72 to
80° during the capture-recapture periods, and up to 86° or so
as a daily maximum. Partly to completely cloudy skies, with
occasional showers, were the rule. Relative humidity ranged
from 85 to 100%.
A small sample of adults from the nearby and largely uncut
rain forest, 8 km. west of Guapiles (by the Rio Toro Amarillo),
was taken; the butterfles were very scarce and scattered there,
despite apparently satisfactory grassy areas along the roads.
GENERAL BIOLOGY & ACTIVITY OF
EUPTYCHIA HERMES
When the first morning sun hits the grassy site of a population
around 7 a.m., the Euptychia begin flying. During the following
two hours, they are quite active and drink sap from crushed
158
T. C. EMMEL
/. Res. Lepid.
sugar cane and suck water from mud in the road bed. By 9:30
a.m., when the full sun is quite intense, there is hardly a butterfly
to be seen in the open. This lack of activity in open areas con-
tinues through the rest of the day. In Guanacaste, it was noted
that a few were flying in the shade or in the undergrowth,
wherever they were protected from the sun and partly protected
from the strong winds. The daily activity cycle is roughly
graphed in Figure 4.
This Euptychia is a low-flying butterfly, clearing the ground
or tops of gress blades by only a few inches. It frequently rests
on the broad-bladed grasses. No oviposition was observed. No
evidence of larvae could be found, but this is usual for satyrid
populations because of the nocturnal larval feeding habits and
the known behavior in several nearctic satyrids of eating the
entire grass blade, leaving little or no evidence of activity.
One mating attempt was observed at 8:45 a.m. February 14 at
Guanacaste. A female landed on a partially-sunlit horizontal
grass blade and a male, which had been following her closely
in flight, landed behind her. He walked rapidly up on her left
side and curved his abdomen around in a U-shape into a copu-
lating position, but the female was skittish and moved away
slightly. At this point, a second male landed ahead of the
female and rushed towards the pair; all three butterflies immedi-
ately flew away in separate directions,
Euptychia hermes was sympatric with three other Euptychia
species at Guanacaste, and with one of the same species at Los
Diamantes in Limon Province.
Analysis of the age composition (as determined by fresh,
intermediate or worn conditions of wings, and the daily addition
of fresh adults) of the male and female samples of all popula-
tions indicated a continuous emergence well before and during
the study period in both the western and eastern populations.
There is no reason to doubt the belief that this Euptychia breeds
continuously throughout the year in these lowland forests (sur-
prisingly even during the dry season in the deciduous forest,
wherever moisture for green grass is available), as larval food
is apparently available at all seasons and the species is known to
occur at all seasons in tropical parts of eastern Mexico ( Emmel,
unpublished data). Therefore the species probably does not
have a diapause stage in these tropical populations. It would
be of interest to carry out comparative physiological and genetic
studies with E. hermes since the extra-tropical populations in
the north, which face severe conditions with a cold winter instead
7(3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
159
of a dry season, apparently have a genetically controlled, obliga-
tory diapause in the larval stage.
A red Orbatid mite was found on the dorsal surface of the
abdomen on each of two females in the Los Diamantes area;
these mites were firmly attached and feeding. No bird attacks
or other predation were observed. No flower-feeding by the
adults was ever noted, so it is unlikely that reduviid bugs, man-
tids or crab spiders are significant predators.
POPULATION SIZE ESTIMATES
1. Guanacaste Results:
During the first three days, 59 adults were marked and re-
leased, with 32 new adults added in sampling on the fourth and
final day of study. Of the 59 releases, 7 (3 males, 4 females)
were recaptured at least once, 1 female was recaptured twice,
and 1 female was recapturd on all thre days following the day
it was marked. No marked butterfly changed from one popula-
tion area to another. Only one individual was recaptured in a
marked population other than E, so estimates of population size
(total number of individuals flying daily) were restricted to
population E, using a simple Lincoln-Index proportion calcula-
tion.
Est. Population Size
Date
110 adults February 14
75 adults February 15
234 adults February 16
Allowing for vagaries of individuals and varying weather con-
ditions, these figures give an approximate population size of
between 75 and 250 adults for a grassy area of only about 30
square meters. It is likely that individuals move back into the
undergrowth in daily wanderings and may be absent from the
grassy stand for a day or two. Variations in apparent flying-adult
population size are known to be due to these environmental and
“wandering” factors, among others (Emmel and Emmel, 1963).
IL Los Diamantes (Limon Province) Results:
A total of 57 adults were marked and released in population
A-B during the first days ofthe study; three more were sampled
on the sixth day. Of the 57 releases, 8 were recaptured once;
none was recaptured twice. The marked irregularity of recap-
ture of marked adults (none on three days) permits estimates
for only the following three dates:
TABLE 1
Sex Ratios in the Guanacaste Populations!
Bailv New Captures (1966)
160
T. C.
EMMEL
/. Res. Lepid.
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7(3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
161
Est. Population Size Date
109 March 6
299 March 8
143 March 9
Since individuals were frequently seen entering and leaving
Area A-B, and since all but one recaptures were females (and
females are notably more sedentary than males in this and all
other butterfly species studied to date ) and none was recaptured
more than once, it seems probable that this supposed ‘popula-
tion” is merely a concentration of individuals in a section of a
much larger population, whose individuals range into the rubber
groves and adjacent fields for at least 30 meters or more (ob-
served distance of flight of several males). This lush grass area
is also likely a favored oviposition site for the females (note
the preponderanee of females here, in Table 2, as compared to
the “normal” preponderance of males in Population C and in
Guanacaste populations ) .
SEX RATIO IN POPULATIONS
The overall sex ratio in the Guanacaste populations was 1.88
males: 1.00 female (Table 1). Most females occured in popu-
lation E here, which apparently was the only stable resident
population in view of the capture-recapture results.
In the Los Diamantes populations, the overall sex ratio was
0.95 males : 1.00 female (Table 2). When the transient “popu-
lation” A-B (sex ratio of 0.58 : 1.00) is considered separately
from the apparently “resident” population G, though, the latter
(sex ratio of 1.41 males to 1.00 females) is seen to be similar
in its male-dominated sex ratio to the Guanacaste populations.
EXTENT OF INTRA- AND INTER-POPULATION
MOVEMENT
There was no observed interchange of individuals between
any of the Guanacaste populations (delineated by shaded areas
on the map in Figure 2); thus these appear to be reproductively
isolated breeding units. The same conclusion is reached for the
Limon populations from the lack of interchange of marked in-
dividuals between Area A-B and Area C at Los Diamantes. From
the available evidence, then, a distance of several score meters
or less ( only 20 meters between areas A and B in Guanacaste )
of unsuitable habitat appears to effectively separate populations
of this Euptychia species. Further study is needed here to de-
Sex Ratios in the Los Diamante s Populations!
162
T, C. EM MEL
/. Res. Lepid.
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7{3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
163
termine the precise amount and type of barrier required for
isolation; quite likely, the barriers to dispersal are intrinsic
(genetically-controlled) as well as environmental, since this
butterfly is capable of flying over thirty meters when pursued
and can fly through tangled undergrowth without much hesi-
tation.
The marked butterfles in subareas Ei and Eg of the Guana-
caste Population E, and in subareas A and B of the Los Dia-
mantes Population A-B, exchanged daily positions back and
forth within the total population area with equal frequencies.
From observation of flight behavior, also, there was no evidence
of territorial or homing behavior in either males or females. This
is in contrast to data obtained on the nearctic lycaenid butterfly,
Plebejus icarioides, where both sexes usually stay in their “home”
part of the population area and will return to it if displaced
(Emmel, ms, in prep.).
DISCUSSION AND CONCLUSIONS
The general findings of the present study concur with many
population parameters characteristic of the majority of investi-
gated temperate-zone butterfles. This Euptychia exhibits about
the same degree of sedentary behavior as the satyrid Cercyonis
oetus (Emmel, 1964, unpublished); the lycaenids Plebejus icar-
ioides (Emmel, ms. in prep.), Philotes sonorensis (Mattoni and
Ralston, ms. in prep.), and Polyommatus icarus (Dowdeswell
et ah, 1940); the nymphalid Euphydryas editha (Ehrlich, 1961,
1965); and the pierid Anthocaris sara (Evans, 1955). In other
words, the species fails to exercise its apparently high degree of
vagility, the ability to cross barriers. Capture-mark-recapture
studies confirmed that while intrapopulational movement occurs
regularly, interpopulational movement is of such insignificance
that these populations are effectively genetic isolates, despite
being separated, in some cases, by only about twenty or thirty
meters of unsuitable habitat.
The species is most active in the early-morning hours and the
later decrease in flight activity may be due to wind (at Guana-
caste) and solar radiation reaching relatively intolerable levels
as the day progresses. Euptychia hermes most likely breeds
continuously throughout the year in these tropical populations,
even in areas having a pronounced dry season. One might sup-
pose that populations increase and disperse to make essentially
continuous huge populations in many areas during the wet
season, when more green grasses would be available. However,
164
T. C. EMMEL
]. Res. Lepid.
the fact that such large continuous populations were not found
in the eastern wet rain forest areas mitigates against this hypothe-
sis. Since the distribution of the species in lowland Costa Rica
seemed closely tied to that of a certain broad-leafed grass (still
to be identified), foodplant specificity may control the butter-
fly’s distribution more than any particular environmental factor.
Such a situation is suspected for the satyrid Cercijonis meacli in
the western U.S., which occurs in widely-scattered, small popu-
lations from 5000-foot-elevation juniper woodlands to 10,000-
foot-elevation mountain pine forests, always in association with
particular grass species (Emmel, unpublished).
SUMMARY
Population sizes, intrapopulation and interpopulation move-
ment, and local and geographic phenetic variation were analyzed
in grassy meadow populations of the satyrid butterfly, Eiiptijchia
hermes Fabricius, located in western and eastern Costa Rica
(Finca Taboga, Guanacaste Province; and Los Diamantes, Limon
Province). Population size was determined by mark-recapture
experiments; a typical population site of 30 square meters in
area had between 75 and 250 flying adults in it during the dry-
season study period.
No movement of marked individuals occurred between popu-
lations separated by as little as 20 meters of dry or non-grassy
areas, yet the butterfies moved freely around within a population
area and when deliberately forced to, could fly more than 30
meters linear distance. Thus, intrinsic (genetic) factors proba-
bly play the major role in limiting flight movement and dispersal.
ACKNOWLEDGMENTS
This research was C4rried out while the author was a partici-
pant in the Organization for Tropical Studies Fundamentals of
Tropical Biology course, February-March 1966, in Costa Rica.
Dr. Charles D. Michener, University of Kansas, kindly reviewed
and commented upon an earlier draft of this manuscript. The
present version is the first of a series of papers analyzing tropical
and temperate population characteristics in Eiiptychia hermes
under the support of a Biomedical Sciences Grant from the
Division of Sponsored Research, University of Florida.
7{3) : 153-165, 1968 (1970)
POPULATION BIOLOGY
165
LITERATURE CITED
DOWDESWELL, W. H., R. A. FISHER, and E. B. Ford, 1940. The
quantitative study of populations in the Lepidoptera. 1. Polyommatus
icarus Rott. Ann. Eugenics, 10: 123-136.
, 1949. The quantitative study of populations in the Lepidoptera.
II. Maniola jurtina L. Heredity, 3i 67-84.
EHRLICH, P. R., 1961. Intrinsic barriers to dispersal in checkerspot but-
terfly. Science, 134 (3472): 108-109.
EHRLICH, P. R. 1965. The population biology of the butterfly, Euphyd-
ryas editha. II. The structure of the Jasper Ridge colony. Evolution,
19: 327-336.
EHRLICH, P. R., and S. E. DAVIDSON, 1961. Techniques for capture-
recapture studies of Lepidoptera populations, /. Lepid. Soc., 14: 227-
229.
EMMEL, T. C., and J. F. EMMEL, 1963. Ecology studies of Rhopalocera
at Donner Pass, California. II. Meteorologic influence on flight activ-
ity. /. Lepid. Soc., 17: 7-20.
EMMEL, T. C., 1964. The ecology and distribution of butterflies in a mon-
tane communitv near Florissant, Colorado. Amer. Midi. Nat., 72: 358-
373.
EVANS, W. H., 1955. Retrieving marked Anthocaris erakirtii. Lep. News,
9: 118.
Journal of Research on the Lepidoptera
7(3) : 166, 1968 (1970)
1160 W Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
A REARING TECHNIQUE FOR SPEEDING
UP THE LARVAL STAGES OF SOME ROOT
OR STEM-RORING LEPIDOPTERA
NOEL McFarland
Entomolofiy Department, S.A. Museum,
Adelaide, South Australia
Larvae that bore in roots, stems, or tree trunks (cossids,
aegeriids, etc.) can sometimes be reared with convenience, and
at a great saving of time and trouble, by providing them with
raw sweet potatoes, white potatoes, yams, beets, turnips, par-
snips, or carrots, etc., into which they can bore.
If they will accept one of these substitutes, growth is often
more rapid than under natural conditions. In Kansas I had excel-
lent results with Frionoxystus robiniae Peck (Cossidae), by pro-
viding the newly-hatched larvae with raw potatoes (both white
and sweet), which they readily accepted, even though in nature
they bore inside cottonwood (Popiilus), and other tree trunks;
growth was completed in less than one year, and healthy adults
emerged shortly after pupation. It is necessary to remove the
larva from the potato, or whatever vegetable is found suitable,
when it has been mostly consumed inside or is beginning to
spoil. Time will vary according to the vegetable used. To start
the larva into a fresh potato, make a hole in the potato and
thrust the larva’s head inside; it will usually proceed to bore in
without further attention. Boring larvae are sometimes inclined
to kill each other when crowded, so they should be permitted to
lead solitary lives, particularly as they grow larger.
166
Journal of Research on the Lepidoptera
7(3) : 167, 1968 (1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
HABITAT: EUPHYDRYAS EDITH A WRIGHTI
Vernal pools are an interesting feature of the coastal mesas
of Southern California. These pools form in shallow depressions
which are underlain by an impervious layer, and are surrounded
by low mounds of earth. These depressions fill with rainwater
in the winter and since there is no drainage, the water remains
until it slowly evaporates in the spring. This novel habitat is
ancient enough to have developed a unique flora and fauna.
Among the plants is Plantago insularis, which in pure stands
resembles a lawn. This is the foodplant of Euphydryas editha
wfighti.
This habitat is being rapidly destroyed as people crowd into
the coastal areas. As a result this butterfly is becoming difficult
to find in areas where it was abundant twenty years ago. Fortu-
nately P. insularis is not confined entirely to vernal pools, sinpe
slow drainage around the lower edges of slabs of rocks, and
perhaps other situations, also provide small areas where this
plant can grow in the foothills. Small colonies of wrighti exist
in some of these spots, hopefully at densities which will permit
survival. (Fig. 1).
Fred Thorne
167
168
THORNE
J. Res. Lepid.
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NOTICES
BOOKS;
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastimil Choc. Paul Hamlyn, Hainlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOCEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McCraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophical
Library, N.Y.
WANTED:
Brephidiurn exilis, B. fea, B. isophtlialma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, California 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the sj^ecies and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systematics, biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
Manuscripts for immediate publication in this JOURNAL. With color
nlay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3x5 card in the form desired and they will be printed in the
next following issue of the JOURNAL.
THE J©UI^N1AL ©F RESEARCH
©NJ THE LEP!J©©FTE^A\
Volume 7 Number 3 September, 1968
IN THIS ISSUE
Population Structure of Oeneis melissa semidea
C. S. Anthony 133
A Hybrid Limenitis from New York
Arthur M. Shapiro and James D. Biggs 149
Population Biology of the Neotropical Satyrid
butterfly, Euptychia hermes.
1. Interpopulation movement, etc.
Thomas C. Emmel 153
Rearing technique for speeding up larval stages
Noel McFarland 166
Habitat: Euphydryas editha wrighti
Fred Thome
167
THE JOURNAL
Number 4
Volume 7
December, 1968
published by
The Lepidoptera Research Foundation, Inc.
at
1160 W, Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz
Associate Editors:
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Florida 32601.
Maria Etcheverry, Centro de Estndios Entomologicos, Casilla 147, Santiago,
Chile.
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Canada.
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by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
cation and the general business office are located at 1160 W. Orange Grove Ave.,
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Journal of Research on the Lepidoptera
7(4):169-181, 1968(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
A STUDY OF
A CONTINUOUSLY BREEDING POPULATION
OF DANAUS PLEXIPPUS
IN SOUTHERN CALIFORNIA COMPARED TO A MIGRATORY
POPULATION AND ITS SIGNIFICANCE
IN THE STUDY OF INSECT MOVEMENT.
F. A. URQUHART, N. R. URQUHART^ and F. MUNGER^
^ Dept, of Zoology and Scarborough College, University of Toronto
^ Dept, of Biology, Whittier College, California
INTRODUCTION
Early in our studies of the movements of the monarch butter-
fly, it was considered that this species in North America formed
a single, gene-flow population the members of which migrated
from north to south in the late summer and fall returning the
following spring ~ thus representing a movement similar in
many respects to that of a species of migratory bird ( Urquhart
1960). However, it was found that in certain parts of North
America there were apparently populations that continued to
breed throughout the entire year, as in southern California and
Florida, or throughout the winter months, as in southwestern
Arizona (Funk 1968). Whether the latter population breeds
throughout the summer months as well is not indicated. It had
also been noted that it is possible to keep breeding populations
throughout the year under laboratory conditions (Urquhart &
Stegner 1966). It was further observed that females of a mi-
grating population, or those found on over-wintering roosting
sites, such as the one located in the Monterey Peninsula of
California, failed to oviposit when brought into the laboratory,
whereas gravid females collected in the field in the summer and
fall laid eggs that gave rise to successive generations (Urquhart
& Stegner op. cit. ). This has made it possible to have a con-
tinuous population for our laboratory experiments on various
aspects of insect physiology.
169
/. Res. Lepid.
170
URQUHART, ET AL
1
7(4); 169-181, 1968(1970) POPULATIONS
171
2-«|.
Fig. 1: Flight lines of the movements of monarch butterflies from
release point at Whittier, California. Explanation in text.
172
URQUHART, ET AL
/. Res. Lepid.
As a result of these considerations,' and in conjunction with
our world-wide studies of the migratory habits of D. plexippus
and with particular reference to the eastern North American
population (Gulf Coast Population, Urquhart 1966) a study of
the movements of the population in southern California (release
point at Whittier) was commenced in 1964.
METHOD
Of the specimens constituting the population studied, most
were reared under natural conditions but some were reared
under conditions of artificial light and temperature. A pre-
liminary study of the data for these two populations indicated
no significant difference in their flight behaviour and therefore
the two populations are considered as a single population reared
and released in Whittier and southern California.
On reaching the imago stage, the specimens were tagged,
using the marginal alar tag method, and released. The time and
place of the relejase of each specimen were recorded. Data con-
cerning direction and duration of flight were obtained from re-
captured specimens that were mailed to our laboratory at the
University of Toronto. All data was recorded and all letters of
correspondence concerning each recapture were kept on file for
future reference. Preliminary maps and flight charts were con-
structed so as to indicate whether or not movements were taking
place and whether or not they were significant.
PRESENTATION OF RESULTS
A total of 8816 imagoes of both sexes were tagged and released
and of these 401 representing both sexes were recaptured and
sent to us. The present study is based on the data from these
recaptured individuals.
Recapture of specimens that had flown more than one mile
from the point of release, and hence indicate a significant move-
ment, were plotted on the map of Los Angeles and Vicinity
(fig. 1). Each release-recovery line represents the flight of one
or more butterflies. The V’ mark indicates the place where the
specimen was recaptured and the number beside the mark shows
how many specimens were recaptured at that particular point.
The flights of specimens which were on the same flight path
were joined together in order to show more clearly the movement
of the population in a particular direction. Names of places
have been included in order to orient the flight patterns and
PERCENTAGE OF SPECIMENS RECAPTURED
7(4);169-181, 1968(1970)
POPULATIONS
173
WEEKLY PERIODS
Fig. 2: Histogram indicating percentage of recaptures at weekly intervals.
174
URQUHART, ET AL
/. Res. Lepid.
hence such place names do not necessarily indicate a particular
point of recapture. Long flights that extended beyond the area
of the map are indicated on the margin with a notation giving
the distance from Whittier to the particular place indicated.
The specimens were released in a well-populated area of
Metropolitan Los Angeles which made it possible to obtain a
high percentage of recaptures — 4.5% as compared to the usual
2% — from all compass points. If releases had been made in a
rural district, then areas of no recaptures would have occurred
owing to the absence of humans and not necessarily an absence
of flights in these particular directions.
An examination of the flight pattern (fig. 1) indicates that
there are relatively few flight lines in the sector extending from
northeast to west and that the majority of flight lines are to the
southwest, south and southeast with the longest flight lines to
the south-southeast. There is a conspicuous lack of flight lines
in the sector southeast to east, with the exception of a few short
flights to the southeast. The absence of long flight lines in this
direction might be due to the presence of the Chino Hills and the
Santa Ana Mountains deflecting the line of flight, or the sparse
human population in this area, or a combination of both factors.
The histogram (fig. 2) shows the time period, in weeks, of
recaptures. As one would expect, when a number of specimens
are marked and released at a particular geographical locality,
there would be more recaptures in the area of release than
remote from it. It wil Ibe noted that a high proportion (20.8%)
of recaptures were made during the first week and had flown
less than one mile. A much smaller number (7.6%) traveled more
than one mile before being recaptured during the first week.
There is a marked drop in the number of recaptures during
the second week, as a result of the thinning out of the popula-
tion away from the point of release. This is followed by a more
gradual decrease in recaptures which finally becomes stabihzed
after the ninth week.
In the fifth and sixth week more specimens are recaptured
after flying more than a mile as compared to those that had
flown less than a mile. A similar situation occurs at week nine
and week fourteen. This indicates that more individuals of the
population had moved away from the point of release than had
remained. There is a marked decrease in the number of speci-
mens recaptured after the tenth week reaching a more constant
level as the tagged population became more thinly spread out
over the countyrside and entered those areas of less dense
human population.
7(4):169~181, 1968(1970)
POPULATIONS
175
Fig, 3: Movement of Monarch butterflies at various times of the year.
Explanation in text.
176
URQUHART, ET AL
/. Res. Lepid.
Thus it would appear from the data here presented that within
this population some individuals tend to be resident (58%) while
others tend to migrate (42%). The proportion of resident indi-
viduals might be considerably less than here indicated, however,
because as a result of a dense population of tagged specimens
located at the point of release, more of them would be recap-
tured before being able to move out. If, for example, some method
could have been employed so that no specimens were recaptured
for an arbitrary period of say two weeks, then there would be
a greater chance for tagged specimens to move out of the con-
gested area thus decreasing the percentage of what appears to
be resident specimens. Of the total number recaptured, 75.5%
were recaptured by the end of four weeks and 89.8% by the end
of eight weeks while only 6.8% were recaptured between the
end of the eighth week and the end of the 20th week. This
can be correlated with the decrease in density as the tagged
specimens spread out over the countryside away from the point
of release.
Fig. 3 presents a graphic analysis of movements of individuals
of the population to compass direction at various times of the
year. Each concentric line represents intervals of four weeks,
commencing at January 1 at the center point. Thus, the first
concentric ring represents January 28; the second ring, Febru-
ary 25; and so on. For ease of reference, the month periods
have been indicated. The direction of flight is given to eight
points of the compass and each recaptured specimen is repre-
sented by a dot. Specimens recaptured at the point of release
have not been included in the chart.
It will be noted that there is a strong southerly movement
between September 9 and December 2. Although movement
tends to be random between January 1 and September 9, there
is a marked tendency for a northeast flow between March 25
and July 15. Very little directional movement is indicated be-
tween July 15 and September 9.
If .we examine the movement for the entire population, with
the exception of those recaptured at the point of release, and
using the method of Williams (1930), in which each arrow point
represents a recaptured specimen it becomes obvious that there
is a very definite movement away from the point of release
towards the southeast and the southwest (fig. 4). There is also
a definite tendency of flights to the northeast.
7(4):169~181, 1968(1970)
POPULATIONS
177
N
NW NE
Fig. 4: Direction of migration of the entire population for the four year
period of investigation. Explanation in text.
178
URQUHART, ET AL
J. Res. Lepid.
CONCLUSIONS
It would appear, as a result of this study and previous ones,
that the monarch butterfly exhibits two types of movement;
one extending over long distances as is the case for the Eastern
North American Population (Urquhart op. cit. ) and others in
which the movement is restricted to shorter distances.
Of those that migrate over long distances, as from the eastern
parts of the United States and eastern Canada to southern
Mexico with a partial return migration, the females enter a
period during which no eggs are produced by the ovaries — an
ovarian dormancy (Urquhart op. cit.). During the autumnal
flight southward such long-distance migrants collect on over-
night roosting sites. On certain over-wintering sites, such as
occur in the Monterey Peninsula of California, they remain
throughout the winter months, (December to February) with no
indication of ovarian activity ( Urquhart op. cit. ) . Under labora-
tory conditions such females taken from the over-night roosting
sites or from the over-wintering roosting sites fail to lay eggs.
In contrast, the offspring from gravid females collected at the
same time as the long-distance migrants were moving south-
ward, continue to lay eggs throughout the winter months. Or,
if females are collected during the early summer and mid-sum-
mer period, eggs will be laid and the offspring continue to do
so throughout the winter months ( Urquhart and Stegner op. cit. ) .
Of those that migrate over short distances, as in the case under
consideration, there is no ovarian dormancy and they do not
cluster on roosting sites. However, as indicated in this study,
the movements do coincide in time with those of the long-
distance migrants of the Eastern North American Population —
eg. a northeasterly trend in the spring and early summer and a
southerly trend in the late summer and fall.
One can hypothesize a definite advantage to an ovarian dor-
mancy for long-distant flights. There would be less delay
caused by oviposition. There would be less weight due to the
absence of eggs. There would be more body fat available for
longer flights. There would be a longer life period, extending
up to six months (September to March) — gravid females on
depositing their full complement of eggs live a much shorter time
and it has been observed in our laboratory colonies that once
oviposition starts, the appendages become brittle resulting in
the loss of tarsi with the resulting destruction of the chemo-
receptors and hence the inability of the individual to locate the
7(4):169-181, 1968(1970)
POPULATIONS
179
source of food or indeed to be stimulated to oviposit; this
process and the relationship between the chemoreceptor mech-
anism and food selection is now being investigated in our labora-
tory.
Ovarian dormancy occurs in over-wintering populations in
northern California and in all of the other states of the United
States and provinces of Canada with the exception of resident
populations in southern California and southern Florida. It is
interesting to note that all of North America, with the excep-
tion of southern Florida and southern California, has diurnal
temperature fluctuations that repeatedly reach freezing or near-
freezing conditions from November through April with marked
fluctuations in late September and October. This variation in
diurnal temperatures can be correlated with the passage of cold
fronts as a result of polar air mass outbreaks from the north-
west (Urquhart op. cit. ). Cold fronts rarely affect southern
California and hence freezing temperatures are not frequently
experienced there. A few examples of average monthly vari-
ations for Los Angeles are as follows: September 75° -59° F.;
October, 72° -55° F.; January and February, 63° -43° F. A
similar situation, with a tendency to higher maxima, occurs in
Florida,
From the above it would appear that ovarian dormancy is
correlated with marked fluctuations in diurnal temperatures in
which the lows repeatedly reach freezing or near-freezing con-
ditions, Such fluctuations are effective in the larval stage pro-
ducing an ovarian dormancy in the adult females. In the ab-
sence of such marked fluctuations ovarian dormancy does not
occur and breeding becomes continuous, assuming the presence
of the host plant. This proposal would also explain the reason
why no ovarian dormancy is experienced in the laboratory
where temperatures are fairly constant thoughout the year
(72° F. ). Similarly, ovarian dormancy was absent in specimens
reared in the greenhouse where light period was the same as
that out-of-doors.
It has been noted that the population under consideration
exhibits movement to the southeast and southwest with a few
direct southerly flights. This peculiarity fits the same pattern
applicable to the Eastern Populaion (Urquhart op. cit.). To
account for this tendency, one may propose the following ex-
planation: That a positive phototaxic response would account
for the southeasterly movement in the a.m. period and south-
westerly movement in the p.m. period with only a slight south
180
URQUHART, ET AL
/. Res. Lepid.
movement during the short meridian period. This concep-
tion is indicated in laboratory observatons in which butter-
flies congregate at the section of the cage where the light is
most intense; thus in the a.m. period they congregate in the
southeast section and in the p.m. period in the southwest
section.
It is probable that many species of insects that are not con-
sidered migrants, although sight observations would indicate
movement, follow a pattern similar to that discussed in this
study. Thus, movements tend to be directional at one time of
the year and not at another, which may occur during the
breeding season for those species which do not possess an ovarian
dormancy period. By utilizing the marginal alar tag system for
the larger species of Lepidoptera it might be found that those
species which are suspected of being long-distant migrants fol-
low a similar pattern. There are periodic migrations over long
distances by various species of Lepidoptera in which one would
suspect a similar pattern on a restricted basis during most years,
but with a definite trend over long distances at other times. Our
studies are now being expanded to include the movements of
other species of Lepidoptera, employing the alar tag system.
SUMMARY
The marginal alar tag system for following the movements
of individuals of a population was employed to find out whether
or not members of a continuously breeding population showed
a tendency to migrate. It was found that a restricted migration
did take place and that the direction and time of movement fol-
lowed the same sequence as the eastern North American popu-
lation that has been shown to travel great distances. The con-
tinually breeding population was compared to one possessing
an ovarian dormancy period. It was suggested that this dor-
mancy period permitted longer flights because of increased
longevity, time saved by not ovipositing, decreased weight and
the availability of stored fatty material. A correlation between
low temperature fluctuations in late summer and fall and ovar-
ian dormancy was indicated. It is suggested that perhaps other
species of insects follow a similar flight pattern to that dis-
cussed in the present study and that in so far as the larger spe-
cies of Lepidoptera are concerned, the use of the marginal alar
tag system might give definitive data upon which an accurate
analysis could be made rather than an analysis based on slight
records. It is further suggested that, although movement on the
7(4):169-181, 1968(1970)
POPULATIONS
181
part of a species may be over a short distance the number of
individuals taking part may vary with seasonal changes, par-
ticularly in the case of unusually long periodic flights.
ACKNOWLEDGMENTS
We are indebted to Mr. J. T. Carlisle of Whittier, California
who was responsible for rearing, tagging and reporting on a
large percentage of the population studied.
This investigation, which is a small part of a much larger
study dealing with the ecology and physiology of Danaus plexip-
pus was made possible by grants to the University of Toronto
from the National Research Council of Canada and the National
Geographic Society.
LITERATURE CITED
FUNK, R. S. 1968. Overwintering of monarch butterflies as a breeding
colony in southwestern Arizona. Journ. Lepid. Soc. 22 ( 1 ) : 63-64.
URQUHART, F. A. 1960. The monarch butterfly. University of Toronto
Press: 361 pp.
URQUHART, F. A. 1966. A study of the migrations of the Gulf Coast
populations of the monarch butterfly (Danaus plexippus L.) in North
America. Ann. ZooL Fenn. 3: 82-87.
URQUHART, F. A. and STEGNER, R. W. 1966. Laboratory techniques
for maintaining cultures of the monarch butterfly. Journ. Res. Lepid. ^
5 (3): 129-136.
WILLIAMS, C. B. 1930. Migrations of butterflies. Oliver and Boyd, Lon-
don, U. K.: 473 pp.
HABITAT — Zerene caesonia eurydice Bdv.
This, the Dog's Head butterfly, is found typically from Men-
docino and Sonoma counties in California into Baja California.
Z. c. caesonia is found from the Great Lakes region to Argen-
tina, in North America from southeastern California to the
Atlantic. Z. c. eurydice should be considered a local offshoot,
or geographic race, isolated along the coast of California where
it occurs closely in connection with its larval foodplant, Amorpha
calif ornica. There is no indication that any other plant will
suffice for long as the larval foodplant.
(continued on page 190)
Journal of Research on the Lepidoptera
7(4):183-189, 1968(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
THE EFFECT OF PTERIN PIGMENTS ON
WING COLORATION OF FOUR SPECIES OF
PIERIDAE (LEPIDOPTERA)
EDWARD J. PFEILER, JR.‘
Humboldt State College, Areata, Calif.
The pterins are a class of chemical compounds with a wide-
spread but sporadic distribution throughout the animal kingdom.
They were first isolated from pierid butterflies where they are
responsible for the orange, yellow and white coloration of the
wings. Although the Pieridae are the only butterflies from which
pterins have been reported, these pigments have been identified
in representatives of other insect orders and in crustaceans.
Pterins have also been reported from certain fish, amphibians,
reptiles and mammals (Fox and Vevers, 1960).
The first major study of pterins was undertaken by F. Gowland
Hopkins (1895), who extracted these white, yellow and red
pigments from various Pieridae. At the time, he did not realize
he was working with an entirely new class of chemical com-
pounds. In fact, he identified the white pigment as uric acid and
stated that the yellow and red pigments were close relatives of
uric acid. By means of more refined chemical tests, it is now
known that the pigments Hopkins extracted were indeed pterins.
The object of this study was to extract and identify the wing
pterins of several species of pierids and to determine if there was
any correlation between wing color and pterin content. The four
species finally chosen for intensive study included the white
Pieris rapae (Linnaeus), the yellow Colias harfordii H, Edwards,
the yellow-orange Colias eurytheme Boisduval, and the orange
Eurema nicippe (Cramer).
1 Current Address: Dept, of Entomology, Washington State Univ,, Pullman, Wash.
183
for pterins.
184
E. J. PFEILER
/. Res. Lepid.
rQ
cd
7(4):183-189, 1968(1970) PTERIN PIGMENTS
185
The pterin pigments from the four wings of a dried specimen
were extracted with 2 ml of 1% ammonia after washing the wings
in about 10 ml of acetone. The pigments were then applied to
Whatman No. 1 chromatography paper, the spots being about
2-3 mm in diameter. The chromatograms were developed by
means of descending paper chromatography using 7:3 propanol/
1% ammonia and 4:1:1 butanol/ acetic acid/ water as solvents.
The solvent front was allowed to advance about 15 cm on the
paper. The chromatograms were then removed from the chroma-
tography tank and dried with a hair drier. The pterin spots were
located under long-wave ultraviolet light and identified by their
characteristic fluorescing colors and Rf values.
Standards were established by chromatographing known
pterins (isoxanthopterin and erythropterin from Aldrich Chemi-
cal Co.; xanthopterin and leucopterin from K and K Laborator-
ies). A small quantity of each standard was dissolved in a few
ml of 1% ammonia and run separately, according to the pro-
cedures outlined above.
RESULTS
In Table I the average Rf values obtained for the pigments of
the four species of Pieridae are compared with those obtained
for the standards. The values represent averages compiled from
the results of a large number of chromatograms. Table II com-
pares the UV fluorescing color with the actual color of the pig-
ments. Using both Rf values and fluorescing colors, it was found
that each species of Pieridae contained the same five pigments,
DISCUSSION
Watt (1964) also characterized the pterin pigments of Colias
eurytheme. Table III compares my results with those of Watt
and in addition shows the Rf values for the standards. All de-
terminations were made using 4:1:1 butanol/acetic acid/ water
and Whatman No. 1 paper. I was unable to obtain a standard
of sepiapterin, but the Rf value and color of fluorescence ob-
served for this unidentified spot agree very closely with the lit-
erature for that of sepiapterin ( Harmsen, 1966; Watt, 1964 ) .
E. J. PFEILER
/. Res. Lepid.
Figures 1-4. 1. Fieris rapae, 2. Colias harfordii, 3. Colias eurytheme, 4. Eu~
rema nicippe. The upper specimen (minus the body) in each figure shows
the typical phenotypic coloration of the species. The lower specimen shows
the effect of removing the wing pterins. In each case the lower specimen
has a pearly lustre which is due entirely to a structural effect. When the
7(4):183-189, 1968(1970)
PTERIN PIGMENTS
187
pterins are removed the scales are not affected and the reflected light
gives the white appearance. The dark color that is found on the wings as
spots and/or rnarginal bands is due to the presence of melanin. The mela-
nin is insoluble in 1% ammonia and therefore is not extracted with the
pterins.
Color of pterins.
188
E. J. PFEILER
J. Res. Lepid.
rn
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7(4): 183-189, 1968(1970)
PTERIN PIGMENTS
189
All four species of Pieridae were shown to contain the same
five pterin pigments but in varying amounts. By assuming a
correlation between strength of fluorescence and amount of pig-
ment present, the relative quantities of each pigment could be
estimated. P. rapae contained large amounts of isoxanthopterin
and leucopterin and smaller amounts of the other three pigments.
C. harfordii, C. eurytheme and E. nicippe contained lesser
amounts of the white pigments and large amounts of the yellow
pigments, especially xanthopterin. The largest amounts of ery-
thropterin were detected in E. nicippe.
It can then be concluded that in this representative sample of
pierids, the whites, yellows, and oranges of the wings are due
to the differential concentration of certain pterins.
LITERATURE CITED
FOX, H. MUNRO, and GWYNNE VEVERS, 1960. The Nature of Animal
Colours. Sidgwick and Jackson Limited, London, 246 pp.
HARMSEN, R., 1966. Identification of Fluorescing and U.V. Absorbing
Substances in Pieris brassicae L. /. Insect Physiol., 12: 23-30.
HOPKINS, F. G., 1895. The Pigments of the Pieridae: a Gontribution to
the Study of Excretory Substances which Function in Ornament. Phil.
Trans. Roy. Soc. of London, B 186: 661-682.
WATT, Ward B., 1964. Pteridine Gomponents of Wing Pigmentation in
the Butterfly Colias eurytheme. Nature, 12: 1326-1327.
HABITAT — Zerene caesonia eurydice Bdv.
( continued from page 182 )
Amorpha calif ornica occurs from sea level to 8000 feet in
elevation. The habitat shown (Fig, 1) is at 6500 feet in the
Mill Creek Canyon, San Bernardino Mountains, California, taken
May 31, 1970. Here Zerene is present in a highly variable
population, mostly in the forms known as eurydice but with a
continuous range of variability to caesonia. Amorpha are the
bushes, ranging from one to four feet high, shown around the
rocks (Fig. 1). A plant is shown in closer view in Fig. 2.
W. Hovanitz
Journal of Research on the Lepidoptera 7(4) : 191-204, 1968(1970)
1160 W. Orange Grove Ave., Arcadia, California, 17. S. A. 91006
© Copyright 1968
HILLTOPPING AS A MATING MEGHANISM TO
AID THE SURVIVAL OF LOW DENSITY SPEGIES
JAMES A. SCOTT
201 Wellman Hall
Univ. of California
Berkeley, California 94720
INTRODUCTION
Hilltopping insects have been reported to be rare by Catts
(1964), Chapman (1954), Dodge & Seago (1954), and Hagen
(1962), but no evidence has appeared for Lepidoptera. The
purpose of this paper is to describe the difference in density of
two groups of butterflies and skippers from Gregory Canyon,
Boulder County, Colorado, those species which show hilltopping
behavior and those without this behavior, and to give possible
explanations why hilltopping species are rare. Mating mechan-
isms other than hilltopping are also discussed. General character-
istics of hilltopping species are given.
HILLTOPPING DEFINED AND EXAMPLES OF
HILLTOPPING
Hilltopping is defined as a behavior of certain insects in which
males fly to the summits of hills and when there remain on the
summit and show perching (“territorial”) behavior or patrolling
behavior, resulting in an unexpected abundance of males on
hilltops. This is a behavioral definition; thus the distribution of
males on hilltops is independent of the distribution of the food-
plant. The foodplant may be only on the hilltop, or it may be a
half mile away. Males may either perch on a shrub or spot of
ground (for instance, Papilio zelicaon. Shields, 1968) or may
“patrol” back and forth on the summit ( Shepard, 1966, for Pieris
occidentalis) . Both types of behavior were noted by MacNeill
(1964) in non-hilltopping situations for males of Hesperia (“oc-
cupation” behavior by H. comma and patrolling by H. lindseyi)^
so it is evident that hilltopping behavior is not fundamentally
different from non-hilltopping behavior; hilltopping behavior
occurs when these activities are transferred to a hilltop. Perching
males may remain on a hilltop for several days; Shields (1968)
191
192
T. A. SCOTT
/. Res. Lepid.
Table 1. Results of six hours of collecting on two hilltops within one
mile of Gregory Canyon (the tops of Flagstaff and Green Mountains),
showing the number of specimens caught and the difference betvjeen
the abundance of these species on the two hilltops sampled and the abun-
dance of the same species in Gregory Canyon, The hilltopping species
increased an average of .91 specimens/hour on hilltops, while the
non-hilltopping species decreased an average of 1.00 specimens/hour
on the hilltops, (*data from Tables 2 & 3) .
A.
B.
Abundance
Abundance
in
Hilltopping species
on
Gregory
male
fern.
hilltops
Can.^
Difference
Erynnis persius
20
3.33
.61
2.72
E, pacuvius
4
.67
.14
.53
E. martialis
4
1
.83
.26
.57
E, afranius
3
.50
.80
- .30
Papilio zelicaon
14
1
2.50
.051
2.45
P, indr a
2
.33
0
.33
P , rutulus
1
1
.33
.16
.17
P. eurymedon
5
.83
.63
.20
Pieris sisymbri
6
1.00
.17
.83
Oeneis chryxus
15
2.50
.24
2.26
Speyeria callippe
9
1.50
.18
1.32
S. edwardsii
4
.67
.12
.55
Vanessa atalanta
1
.17
0
.17
means
1.17
.26
.91
Non-hill topping species
Coenonympha tullia
1
.17
2.47
-2.30
Oeneis uhleri
2
.33
1.81
-1.48
Callophrys apama
2
.33
1.91
-1.58
C. polios
2
.33
1.26
- .93
Celastrina argiolus
7
9
2.67
1.40
1.27
means
.77
1.77
-1.00
7(4):191~204, 1968(1970) MATING MECHANISM
193
recaptured marked P. zelicaon up to a month after release. Some
species may visit hilltops only once, however, since of 46
Vanessa cardui males that Shields released on a summit, none
were recaptured after more than a day. Hilltopping Pieris
occidentalis do not stay long in any area (Shepard, 1966).
A ‘'hilltopping species” is defined as a species which has been
observed to show hilltopping behavior. Likewise, a non-hill-
topping species is a species which does not show hilltopping
behavior in the localities the author has studied. A rigid black-
and-white separation of butterflies into hilltopping and non-
hilltopping species is somewhat artificial; for some species of
Hesperia, Erynnis, and Papilio zelicaon, hilltops probably serve
as the primary site of mating, but for other species, such as
Speyeria and Ochlodes sylvanoides, hilltops are probably minor.
Ochlodes sylvanoides males perch on bushes in clearings both
on hilltops and on flat areas. Nevertheless, this separation is
presently justifiable until more is known about each species.
Papilio zelicaon will serve as an example of a hilltopping species.
In the spring of 1966 only one male and two females were
caught in Gregory Canyon during almost sixty hours of collect-
ing, but in one short trip to the top of Green Mountain (about
one-half mile from Gregory Canyon ) eight male P. zelicaon were
caught in less than two hours. The males fly in rather fixed paths
around the rock and through the trees on the summit; if missed,
specimens usually return a few minutes later. Table 1 show the
results of a brief period of collection on two hilltops near
Gregory Canyon. The proportion of hilltopping species present,
13 out of a total of 18, was 72%, whereas it was 42% in Gregory
Canyon where there are no hilltops. The densities of hilltopping
species and nonhilltopping species were 1.17 and .77 respectively
on the hilltops, while the densities for the same species in
Gregory Canyon were .26 and 1.77.
GREGORY CANYON
During the spring of 1965, 1966, and 1967, the author made
extensive collections of butterflies and skippers in Gregory Can-
yon, Boulder County, Colorado, a small foothills canyon on the
eastern slope of the Front Range. It is less than a mile in length.
The southern wall is covered with dense Douglas fir forest which
is for the most part unsuitable for butterfly flight. The south-
facing side of the canyon is a grassy slope with scattered pon-
derosa pines. In the bottom of the canyon is a variety of riparian
shrubs and trees. Extensive collecting was done on the bottom
and south-facing slope of the canyon, on the eastern slope of
194
J. A. SCOTT
/. Res. Lepid.
Table 2. The abundance of nonhilltopping butterflies and skippers in
Gregory Canyon.
Dates of
Specimens
Species
Capture s
per hour
Ambylysclrtes vialus (Edwards)
4-v to 30-v
.20
A. aenus Edwards
16-v to 30-v
.15
A. oslari (Skinner)
13-v to 30-v
2.06
Euphyes vestris (Bolsduval)
28-v
4.00XX
Poanes taxiles (Edwards)
30-v
.33XX
Polite s themistocles (Latreille)
29-v to 30-v
1.40
P. mystic dacotah (Edwards)
29-v
.14XX
Oarisaa garita (Reakirt)
24-v to 30-v
3.37
Pholisora catullus (Fabricius)
16-v to 24-v
.39
Pyrffus mralis (Bolsduval)
2-v to 13-v
.11
Epargyreus clarus (Cramer)
19-v to 30-v
.33
Papilio multicaudata Kirby
5-v to 29-v
.18
Colias alexandra Edwards
29-v
.14XX
C. philodice Godart
27-lii to 24-v
.38
Antho carls sara julla Edwards
14-v to 30-v
.10
Coenonympha tullia ochracea Edwards
5-v to 30-v
2.47
Oeneis uhleri (Reakirt)
26-iv to 23-v
1.81
Euptoieta claudia (Cramer)
29-v
.14X
Phyclodes campestrls camlllus Edwards
13-v to 30-v
.62
P . tharo s ■ ( Drury)
13-v to 30-v
.41
P. pallida (Edwards)
19-v to 29-v
.62
Nymphalis antiopa (Linnaeus)
13-iii to 27-lli
.76
Polygonia zephyrus (Edwards)
13-111 to 23-v
.67
P. satyrus (Edwards)
14-iv to 28-iv
.11
Limenitis wiedemeyeri Edwards
29-v
.14XX
Callophrys polios (Cook & Watson)
27-iil to 23-v
1.26
C. erephon (Bolsduval)
30-iii to 24-v
1.12
C. fotis schryveri (Cross)
30-iii to 5-v
.40
C. apama homoperplexa Barnes & Benjamin
14-iv to 30-v
1.91
C. sheridanii (Edwards)
26-lli to 15-v
.81
Plebejus mellssa (Edwards)
5“V to 30-v
.40
P. acmon lutzi dos Passes
13-v to 23-v
.071
P. icarioides lycea (Edwards)
14-v to 30-v
4.24
Glaucopsyche lygdamus oro Scudder
15-iv to 29“iv
2.74
Scolltantides piasus daunia (Edwards)
14-v to 30-v
1.36
Eve res comyntas valeriae Clench
2-v to 30-v
.56
Philo tes enoptes ancllla Barnes & McD.
5-v to 30-v
.68
Celastrina arglolus cinerea (Edwards)
14-iv to 30-v
1.40
mean
1.034
XX-data thrown out because less than
s =
1.028
ten hours of collecting
7(4):l91-204, 1968(1970) MATING MECHANISM
195
Flagstaff Mountain and on the north slope of Chautauqua Mesa,
both grassy hillsides with a variety of herbs, at the mouth of the
canyon. Brief collecting results on the tops of two nearby hills,
Flagstaff Mountain and Green Mountain are compared with
results in Gregory Canyon.
METHODS
To find out if hilltopping is more prevalent among the rarer
species, the densities of butterfly species in the localities outlined
above were studied. Average density is defined in this study as
the total number of individuals of each species divided by the
total number of hours spent collecting in the area during the
flight period of the species under consideration. For instance,
Amhlyscirtes oslari was collected in Gregory Can. from May (v)
13 to May 30. Since 110 specimens were taken in the canyon
during this time span, and a total of 53.5 hours were spent
collecting in the canyon on and between May 13 and 30, the
density is 110/53.5 or 2.06 spec. /hour. It can be seen that this
abundance value measures neither total population size nor
density, but is an estimate of average density over the total area
during the flight period. Since the flight periods of many species
were much greater than those of other species, the ratio of
specimens per hour was preferred to the number of specimens
as an indicator of relative density since it measures the average
density of a particular species during the flight period sampled.
However, in the statistical test below, both methods of measur-
ing the density were used, and produced similar results. The
densities for each species are in Tables 2 and 3. Data for species
with less than 10 hours of collecting are unreliable and were not
used in the statistical tests. Species with less than 10 hours of
collecting were usually those just beginning their flight period
at the end of May; thus the number of specimens collected may
not represent the abundance later on in the flight period.
Several criticisms of this type of method for sampling the
density are pertinent. First, it is very crude. It would be best to
have sampling programs for each species or run mark-release
studies for each species to estimate the population size. These
studies are impossible for the large number of species considered,
and it is necessary to study large numbers of species to minimize
the effect of “abnormal” species in making generalizations about
hilltopping and nonhilltopping species. Second, the collector
could be prejudiced in sampling. An attempt was made to collect
everything flying. Since the species likely to be undercollected
are the common species in most instances, prejudiced collecting
196
J. A. SCOTT
/. Res. Lepid.
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Figure 1. Abundance of butterflies in Gregory Canyon, 1966.
7(4):191-204, 1968(1970) MATING MECHANISM
197
would shift the results so that the less abundant species would
appear to be more common. The results actually obtained, that
the hilltopping species are less abundant, are thus not affected.
An attempt was also made to collect the area uniformly. The area
was sampled by walking across the lower slopes of Chautauqua
Mesa to the southeastern slope of Flagstaff Mountain, and then
proceeding up Gregory Canyon along a trail near the stream
bottom. The south-facing slope of the canyon was traversed, and
the route then reversed. Varying amounts of time were spent
each day depending upon weather conditions, etc. Collecting was
done between 9 a.m and 5 p.m., usually between 10 and 4. A
third criticism is whether the year or the locality is unusual,
resulting in abnormal abundance of certain species. Every local-
ity of course varies in the abundance of certain species, but
since the number of both hilltopping and non-hilltopping species
is fairly large, the average abundance figures of the two groups
are assumed to be not unduly affected by abnormal abundance
values.
ABUNDANCE OF HILLTOPPING VERSUS
NONHILLTOPPING SPECIES
The results are shown in Tables 2 and 3 and are graphed in
Figure 1. 2023 specimens were collected. The average values for
the number of specimens per hour were found to be .28 for the
hilltopping species and 1.03 for the nonhilltopping species. Thus
the hilltopping species are one fourth as dense on the average
as the nonhilltopping species. The variations in density within
each group was great, with standard deviations of .23 and 1.03
for hilltopping and nonhilltopping groups respectively, and the
group_s overlapped considerably. The second lowest density was
recorded for Plebejus acmon, a non-hilltopping species, and the
fifteenth highest density was recorded for Erynnis lucilius, a
hilltopping species. But in only 6 of 21 species of hilltopping
butterflies was the value above .26, and in the non-hilltopping
species only 7 of 32 had values .32 or below. A test of the
hypothesis that there is no difference in density of the hilltopping
and non-hilltopping groups is shown in Table 4. The probability
that the results obtained are due to chance is .002 (Fishers exact
method). Another test of the hypothesis that there is no differ-
ence in the absolute number of specimens caught of hilltoppers
and non-hilltoppers resulted in a probability of .008 that there is
no difference.
198
J. A. SCOTT
/. Res. Lepid.
Table 3. The abundance of hilltopping butterflies and skippers in
Gregory Canyon, 1966.
Dates of
Specimens
Species
Capture s
per hour
Hesperia juba (Scudder)
19-v to 29-v
.18
Erynnis persius frederickl Freeman
28-iv to 30-v
.61
E. pacuvius (Lintner)
1-v to 29-v
.14
E. martialis (Scudder)
1-v to 30-v
.26
E. icelus (Scudder & Burgess)
3-v to 24-v
.26
E. lucilius afranius (lintner)
27-v to 30-v
.80
E. telemachus Burns
3-v
.25XX
Thorybes mexicana nevada Scudder
25-v to 30-v
.093
T. pylades (Scudder)
14-v to 30-v
.46
Papilio zelicaon Lucas
29-iv to 28-v
.051
P. indra Reakirt
25-v to 28-v
o**xx
P. rutulus Lucas
28-v to 30-v
.16
P. eurymedon Lucas
19-v to 29-v
.63
Euchloe Olympia rosa (Edwards)
1-v to 13-v
.083
E. ausonides coloradensis (H. Edwards)
14-iv to 30-v
.63
Pierls sisymbri elivata Barnes & Ben j .
14-iv to 28-v
.17
Oeneis chryxus (Doubleday)
23-v to 30-v
.24
Speyeria callippe meadi (Edwards)
24-v to 28-v
.18
S. edwardsii (Reakirt)
25-v to 30-v
.12
S. coronis halcyone (Edwsirds)
29-v
.14XX
Poladryas arachne (Edwards)
27-v
.25XX
Chlosyne ismeria car lota (Reakirt)
24-v to 27-v
.14XX
Nymphalis milberti furcillata (Say)
13-iii to 2-v
.12
Vanessa cardui (Linnaeus)
15-iv to 15-v
.075
V. atalanta (Linnaeus)
28-v
0**XX
Strymon melinus franki Field
.17XX
Callophrys spinetorum Edwards
3-v to 13-v
.13
C. augustinus irioides (Boisduval)
26-iii to 4-v
.60
XX- data thrown out because less than ten
hoiirs collecting
mean .285
S = .231
specimens seen but could not be collected
7(4): 191-204, 1968(1970) MATING MECHANISM
199
POSSIBLE EXPLANATIONS FOR THE LOWER
DENSITY OF HILLTOPPING SPECIES
Fewer specimens of hilltopping species could have been
caught in the canyon because the males were on nearby hilltops.
However, the percentage of males of 1957 total specimens for
which the sex was determined was 74,2% (with 95% confidence
limits of 53% and 86% ) for hilltopping species and 82.6% (with
95% confidence limits of 66% and 96%) for nonhilltopping
species. The greater number of males is not due to female mor-
tality or actual skewed sex-ratios in the population at birth,
since rearing butterflies usually results in equal numbers of
males and females (Shields, 1968), but is probably due to more
overt behavior on the part of the males; females are inconspicu-
ous due to oviposition behavior. Therefore, the disparity in
abundance is not explained by different sex-ratios or by the
absence of males due to their flying to distant hilltops. Also,
the two hilltops nearest to Gregory Canyon are relatively far
from the collecting area, are 600 to 1500 feet higher than
Gregory Canyon, and are comparatively poor in number of
species and individuals present.
A hypothesis of the ecological function of hilltopping suggests
that male butterflies are on hilltops for mating purposes. This
theory suggests that males, which in butterflies usually emerge
earlier than females, visually orient and fly to the hilltops and
that the females, when they emerge, then fly to hilltops, mating
occurs, and then the female leaves to lay her eggs. Presumably
if the female meets a male before she arrives at a hilltop, mating
would occur and she would then not visit a hilltop. This theory
is supported by observations of mating pairs on hilltops, a high
percentage of virgin females of hilltopping species on hilltops,
and ah experiment in which virgin female Papilio zelicaon flew
to hilltops (Shields, 1968). However, no studies have been con-
ducted which compare the number of matings [or percent vir-
gins] on a hilltop with the number [or percent] off a hilltop.
This theory suggests several reasons why hilltopping species
should be less dense than non-hilltopping species.
Hilltopping may provide a rendezvous for very rare species
which otherwise may not produce enough individuals to survive.
In general the number of contacts between individuals should
be proportional to the density of the population; in low-density
populations a hilltop rendezvous may increase contacts to the
extent that the percent of females which successfully mate is not
200
J. A. SCOTT
/. Res. Lepid.
Table 4» Contingency tables of the abundance of the species.
Figures above slash refer to niimber of species above or below mean
(.737 specimens per hour) using the number of specimens per hour as
an indicator of abundance. Figures below slash refer to number of
species above or below mean (37.96 specimens) using the number of
specimens caught as an indicator of abundance. Probabilities that
tables as or more extreme than the ones above occur by chance alone
are .002 using number of specimens per hour, and .008 using total
number of specimens (Fisher’s exact method).
No. of species
No, of species
with abundance
with abundance
Hilltopping
Species
greater than mean*
1/
/2
less than mean*
20/
/19
Totals
21
Non-hilltopping
Species
14/
/15
18/
/17
32
15/
/II
38/
/36
Totals
53
7(4):191-204, 1968(1970) MATING MECHANISM
201
seriously reduced. A nonhilltopping species in a marginal locality
where it is rare may not survive. This may be the reason why
most hilltopping species are widespread and less dense, which
many nonhilltopping species are local and dense.
Hilltopping can be effective only for low density species,
because 1) at high densities on hilltops interference between
males prevents mating with females, and 2) the number and
area of hilltops is limited. Point 1) is based on the author’s and
Shields’ (1968, pp. 134, 139) observations. Point 2) is supported
below. If a species is common, only a small proportion of the
males can occupy a hilltop, so that most males will be forced
into nonhilltop situations.. As population density rises, the proba-
bility that a female will meet a male before reaching a hilltop
therefore increases, so that hilltopping is less important for
commoner species. The few males on hilltops could not possibly
inseminate all the females in a common species, so that most
matings will occur with males which remain at the breeding site
or which are between the breeding site and the hilltop. Because
hilltopping is less useful for common species, selection should
eliminate the hilltopping response since males which remain at
the breeding sites will contribute more genes to the next
generation.
It is possible that hilltopping could act as a “population con-
trol” agent to prevent the population from exceeding a certain
density. However, the author’s preliminary data for Hesperia
pahaska and Amblyscirtes simiiis indicates that hilltopping breaks
down when these species are dense and mating occurs on hill-
sides and sloping ridges as well as hilltops.
Hilltopping may centralize the gene pool of a population. It is
conceivable that this could reduce the density of the species in
an area by preventing microenvironmental adaptation. This
would be difficult to prove, and two different microhabitats
within the dispersal range of a species may not exist .
CHARACTERISTICS OF HILLTOPPING SPECIES
Hilltopping species are in general large, fast-flying, solitary
species with more widely scattered and less abundant foodplants
than non-hilltopping species, which tend to be small, weak-flying,
colonial species with common or clumped foodplants. The aver-
age size of hilltopping species is 22.6 mm in Gregory Ganyon,
while the size of nonhilltopping species is 17.5 mm (based on
the average of the left front wing length of three males); this
difference is significant at the 5% level. The average abundance
202
]. A. SCOTT
/. Res. Lepid.
of the host plant is 1.89 for hilltopping butterflies, and 2.68 for
non-hilltopping butterflies (based on a rating system from 1-rare
to 5-abundant of known Gregory Canyon foodplants by the
author); this difference is significant at the 1% level, although
admittedly the system for rating the plant is imprecise. All of
the hflltopping species in Gregory Canyon are “fast-flyers” with
the exception of Poladryas arachne, whereas less than half of the
species of non-hilltopping species fit into this subjective category.
A hilltopping species must be highly motile, of course, to reach
distant hilltops. Hilltopping is probably more prevalent in dry or
sparsely vegetated mountainous areas, since more small hilltops
occur in eroded foothill and chaparral regions than in flat areas,
and few hilltopping males occur on densely forested hilltops.
OTHER MATING MECHANISMS
There are methods other than hilltopping of bringing the sexes
together from a distance. These other mechanisms put hilltop-
ping in proper perspective and are listed below. A fundamental
difference between these mechanisms and hilltopping is that
many males can participate in the following mechanisms whereas
fewer males can fit on a hilltop; therefore the following mechan-
isms are operable at much higher densities. They may not be
mutually exclusive.
1 ) Chemoreception is known to be very important in the long-
distance location of females by males in moths, and in the court-
ship of moths and butterflies (Jacobsen, 1965). It may prove to
be important in location of females by males in Heliconius
(Edwards, 1881; Bellinger, 1954) and Parnassius. For most
butterflies, however, the maximum distance of attraction is
limited by sight, while chemoreception is important only within
a few meters of the female by the release of pheromones from
hair pencils, androconial scales, etc.
2) Foodplant congregation. Most butterflies, especially the
weak-fliers, spend their entire lives, except for brief forays in
search of mud or flowers for nourishment, around stands of the
foodplant, and therefore have a built-in mechanism for bringing
the sexes together. Often both sexes are limited both to food-
plant and to certain areas of the environment such as rockslides
(Erebia magdalena), bogs (Boloria frigga), or freshwater springs
(Speyeria nokomis), which may or may not be the only locations
of the foodplant. The behavior of these species usually limits
them to these areas so that mating is possible with “random”
flight by both sexes or by patrolling of the area by males as in
7(4hmi-204, 1968(1970) MATING MECHANISM
203
Boloria (J. Shepard, unpublished ) .
3) In some species the males occupy small areas along the
bottom of a gully or canyon, presumably for mating purposes.
Males may occupy an area for some time, but this behavior may
not be territorial since the males may wander to another gully
and show the same behavior. Butterflies which show this be-
havior are Polygonia, Euptychia, Callophrys apama, Papilio
rutulus, and Amblyscirtes oslari. The author conducted mark-
release studies in Gregory Canyon in 1967. 40 C. apama males
were marked and released in a small gully, and of these 13 were
recaptured, including three which were recaptured twice each,
and individuals which were recaptured at the same spot after
2, 5, 5, 9, 19, and 19 days. 46 male Polygonia zephyrus were
released, and 7 were recaptured, including males which were
recaptured after 1, 8, 9, 9, 10, 13 days. One individual moved a
distance of 1000 feet and then was recaptured, then moved a
distance of 200 feet before being recaptured again; all others
were recaptured near the place of release. Amblyscirtes oslari
males show this type of behavior in roadside ditches and small
gullies.
SUMMARY
Hilltopping species appear to be a heterogeneous ■ taxonomic
and behavioral assemblage. They are characterized by the trans-
ference of mating behavior to hilltop situations. The behavior
of Papilio zelicaon and Hesperia pahmka, in which the males
perch on hilltops, is similar to the behavior of Polygonia zephyrus
and Amblyscirtes oslari, non-hilltopping species in which the
males perch in gullies. By contrast, the behavior of Pieris oc-
cidentalk, a hilltopping species in which males patrol the sum-
mit^ is quite different. A separation of butterfly species into
two groups, namely, those in which males perch, and those in
which males continually fly in search of females (patrol), is a
more widely applicable classification of their mating behavior.
Hilltopping species have many traits in common, however.
They do not congregate about the foodplant but instead tend
to be large, strong-flying, solitary species with more widely
scattered foodplants than other species. Populations of hill-
topping species are less dense than those of other species since
selection favors the development of hilltopping in low density
species, but hilltopping confers little or no advantage to species
with high numerical density. Common species would benefit
from hilltopping only when their populations fall to low levels.
204
]. A. SCOTT
/. Res. Lepid.
Hilltopping may be selected for at low population levels, and
remaining near the foodplant may be selected for at high levels,
so that the advantage of hilltopping for a particular species
depends on its average density and the fluctuations from this
average.
ACKNOWLEGMENTS
The author wishes to thank J. A. Powell and J. H. Shepard for
critically reading the manuscript.
LITERATURE CITED
BELLINGER, PETER F. 1954. Attraction of zebra males by female pupae.
Journ. Lepid. Soc. 8:102.
CATTS, E. P. 1964. Field behavior of adult Cephenemyia. Canad. Entomol.
96:579-585.
CHAPMAN, J. A. 1954. Studies on summit frequenting insects in western
Montana. Ecology 35:41-49.
DODGE, H. R., & J. M. SEAGO, 1954. Sarcophagidae and other diptera
taken by trap and net on Georgia mountain summits in 1952. Ecology
35:50-59
EDWARDS, W. H. 1881. On certain habits of Heliconius charitonius L.,
a species of butterfly found in Florida. Papilio 1:209-215.
HAGEN, K. S. 1962. Biology and ecology of predaceous Coccinellidae.
Ann. Rev. Entomol. 7:289-326.
JACOBSEN, M. 1965. Insect sex attractants. John Wiley, New York.
154 pp.
MacNEILL, C. D. 1964. The skippers of the genus Hesperia in western
North America. Univ. Calif. Publ. Entomol 35:1-221.
SHEPARD, J. H. 1966 A study of the hilltopping behavior of Pieris occi-
dentalis. Pan-Pacific Entomol. 42:287-294.
SHIELDS, A. O. 1968. Summit congregation behavior of butterflies on a
southern California hill. Journ. Res. Lepid. 6:69-178.
NOTICES
BOOKS:
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastimil Choc. Paul Hamlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophieal
Library, N.Y.
WANTED:
Brephidium exilis, B. fea, B. isophthalma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, California 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systematics, biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
Manuscripts for immediate publieation in this JOURNAL. With color
irtay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3x5 card in the form desired and they will be printed in the
next following is.siie of the JOURNAL.
THE J0UIRNJAL ©F RESEARCH
©Mi THE LEFIJ©©RTERA\
Volume 7 Number 4 December, 1968
IN THIS ISSUE
Population of Danaus plexippus
in Southern Calfornia
F. A. Urquhart, N. R. Urquhart, and F. Munger 169
Habitat: Zerene caesonia eurydice
W. Hovanitz 182
The Effect of Pterin Pigments on Wing
Coloration of Four Species of Pieridae
Edward J. Pfeiler, Jr. 183
Hilltopping as a Mating Mechanism
to Aid the Survival of Low Density Species
James A. Scott
191
published by
The Lepidoptera Research Foundation, Inc.
at
1160 W. Orange Grove Ave., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz
Associate Editors:
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Canada.
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Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.
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THE JOURNAL OF RESEARCH ON THE LEPIDOPTERA is published four times a
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by THE LEPIDOPTERA RESEARCH FOUNDATION, INC. The office of the publi-
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THE LEPIDOPTERA RESEARCH FOUNDATION, INC. THE LEPIDOPTERA
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Second C^lass postage paid at Arcadia, California, U.S.A.
8(1):1-15, 1969(1970)
Journal of Research on the Lepidoptera
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
SCANNING ELECTRON MICROSCOPY
ON WING SCALES OF COLIAS EURYTHEME
JOHN M. KOLYER and ANNEMARIE REIMSCHUESSEL
55 Chimney Ridge Drive, Convent Station, New Jersey, U.S.A.
Optical microscopy discloses that the scales on the wings of
Lepidoptera may be ribbed lengthwise, with perpendicular cross-
ribs to give a network ( Gentil, 1935 ) , but finer details cannot be
resolved. Transmission electron microscopy has been utilized
to study the fine structure of Morpho scales (Gentil, 1942; Kin-
der and Suffer!, 1943; Richards, 1944), whose iridescent colors
are “structural” and result from diffraction of light by ridges on
the scale rather than from the present of pigments. However,
in the family Pieridae, including the genus Colias, the yellow
and/or orange colors are not structural, and transmission elec-
tron microscopy has been reported to disclose round and spindle-
shaped aggregations of pigment (Yagi, 1954), which consists
of a number of pteridine compounds (Watt, 1964). The black
scales in the border are colored by melanins.
The present work was undertaken on the premise that the
recently-developed method of scanning electron microscopy
(SEM) should be particularly well-suited, due to its advan-
tageous magnification range (45-30,000 X) and depth of focus,
to examination of the surface structure of the scales. Colias eury-
theme (Boisduval) was chosen as an example. A particular ob-
ject was to note possible variations in the fine structure of scales
from different areas of the wing.
METHODS
Figure 1 shows a specimen, male, with indication of the areas
examined. Small portions of these areas were cut out with a
scalpel, and each was mounted on a specimen stub. The speci-
mens were vapor-coated with a thin (300 Angstroms) layer of
gold/palladium alloy (60/40) to render them conductive, a pre-
requisite for examination by SEM. The SEM instrument was
a JEOLCO JSM-2, operated at an accelerating voltage of 25 kv.
Photomicrographs were prepared with Polaroid P/N Type 55
film at a scan speed of 50 seconds per frame.
1
iMITHSONIAK
iMSTITUT-OK
2
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Fig. 1. — Colias eunjtheme, male. Right forewing, showing location of scales
examined.
WING SCALES
3
'8(1):1-15, 1969(1970)
Fig. 2a.— Upper surface, orange scales.
To expose the underside of the scales, the wing was pressed
onto a surface coated with contact adhesive (the backing used
for Polaroid color prints), and the wing membrane was peeled
off to leave the scales perfectly transferred. This method is
successful because the peduncles (stems) are loosely held in
sockets on the membrane, the scales of the upper and under
layers being attached at alternating sockets ( Gray, 1961 ) .
4
KOLYER AND REIMSCHUESSEL
J. Res. Lepid.
Figs. 2b, c. — Upper surface, orange scales.
8(1):1-15, 1969(1970)
WING SCALES
5
Fig, 2d.~Upper surface, orange scales.
OBSERVATIONS
Results are shown in Figs. 2-7. It is interesting that there are
marked differences in fine structure among the four varieties of
scales whose upper surfaces were examined and also between
the upper and lower surfaces of the same (orange) scale.
Butterfly scales long have been described as hollow, as sug-
gested by the holes visible in the photomicrographs. The hollow-
ness of the peduncle seems apparent in Figs. 4b and c. It has
been speculated ( Fortier, 1932 ) that the scales and peduncles of
the genus Parnassius are hollow and therefore admit air, com-
municate with tracheal capillaries in the wing, and play a role
in respiration.
6
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Fig. 3a. — Upper surface, washed orange scales.
The fine structure of the orange scales visible at 10,000X
(Figs. 2b and c, 3b and c) resemble a “double grating” or net-
work. The distance between the lengthwise ribs is approxi-
mately 1.5 microns, and that between the cross-ribs or connect-
ing ribs ranges from approximately 0.5 to 0.7 micron. The thick-
ness of the cross-ribs is approximately 0.07 micron. In some
orange scales the cross-ribs appear to be partially interconnected
by a thin skin or membrane (Fig. 2b) whereas in other orange
scales most of the cross-ribs are not interconnected but ex-
hibit small ellipsoidal structures that appear to be suspended
from them (Figs. 2c and d). The above-described two types of
orange scales are found in different positions with respect to
the “shingling” arrangement (Fig. 2a) on the wing membrane;
the scales with the ellipsoidal particles (Fig. 2c) occupy the
lower layer and are partially covered by the upper-layer scales
shown in Fig. 2b.
8(1):1--15, 1969(1970)
WING SCALES
7
Figs. 3b, c. — Upper surface, washed orange scales.
8
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Fig. 4a. — Under surface, orange scales.
The orange color was removed completely by dipping a wing
first in 95% ethanol and then for only 20 seconds in 20%
aqueous ammonia (the color was transferred to the solution
as the pteridines were dissolved as their ammonium salts).
Then the wing was dipped in water, then ethanol, and allowed
to dry in the air. Photomicrographs of the washed scales are
shown in Fig. 3. The treatment seemed to make no change in
the upper-layer scales (Fig. 3b vs. Fig. 2b), but in the case
of the under-layer scales the suspended particles appear to
have been largely removed to give a more open network (Fig.
3c vs. Fig. 2c). Whether and to what extent the ellipsoidal
particles are related to the color remains to be established.
The under surface of the orange scale shown in Fig. 4 ap-
pears to be without much detailed fine structure. There are no
ribs except on the peduncle and the periphery of the scale.
This observation suggests that the scale resembles a hollow
pouch consisting of two significantly different sheets— a con-
tinuous bottom membrane and a cross-ribbed upper sheet which
is more or less porous depending on the type of scale and its
position on the wing.
8(1):1-15, 1969(1970)
WING SCALES
9
Figs. 4b, c.~Under surface, orange scales,
10
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Fig. 5a. — Upper surface, black scales.
The fine structure of the upper surface of the black scales
(Fig. 5) is strikingly different from that of the orange scales.
The distance between the lengthwise ribs is approximately 3
to 6 microns as compared to 1.5 microns in the orange scales.
Also, the trough-like material between the lengthwise ribs of
the black scales displays intricate patterns which cannot be
described as “cross-ribs” (Figs. 5b and c).
Interspersed among the black scales are brightly-colored
yellow scales in which the distance between the lengthwise
ribs is approximately 3 to 4 microns. The presence of cross-ribs,
and particulate matter in some areas, is indicated (Fig. 6b).
8(l)tl~15, 1969(1970)
WING SCALES
11
Figs. 5b, c. — Upper surface, black scales.
12
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Fig. 6a, b, — Upper surface, yellow scales.
8(1):1-15, 1969(1970)
WING SCALES
13
The final type of scales examined, the pink fringe scales, ex-
hibit lengthwise ribs that are approximately 2 to 4 microns
apart; the inter-rib distance varies from a minimum of about
2 microns at the basal region to a maximum of about 4 microns
toward the tip of the scale. Tilting of the specimen showed
clearly that the lengthwise ribs are composed of overlapping
short narrow ‘"scales” (Fig.' 7c). The material connecting the
lengthwise ribs in this case forms a continuous trough and ap-
pears to be supported by faintly- visible cross-ribs.
14
KOLYER AND REIMSCHUESSEL
/. Res. Lepid.
Figs. 7b, c. — Upper surface, pink fringe scales.
8(1): 1-15, 1969(1970)
WING SCALES
15
SUMMARY
Fine structure varied greatly with color and position. The
upper surface of an orange scale, cross-ribbed and perforated
between the lengthwise ribs (1.5 microns apart), was strikingly
different from the smooth and continuous lower surface as well
as from the upper surface of a black scale, on which the ribs
(5 to 6 microns apart) were connected by intricately-patterned
“troughs”. The peduncles (stems), as well as the scales them-
selves, appear hollow.
ACKNOWLEDGMENTS
We thank the Corporate Chemical Research Laboratory of
the Allied Chemical Corporation for kindly providing facilities
for this work and Mr. Ronald Galante for his assistance in pre-
paring the photomicrographs.
LITERATURE CITED
GENTIL, K. (1935). Der Bau der Schillerschuppen von Papilio paris. En-
tomologishe Rundshau, 52:230-232.
(1942). Elektronmikroskopische Untersuchung des Feinbaues
schillernder Leisten von Morpho-Schuppen. Zeitschrift fur Morphologie
und Okologie der Tiere, 38(2) : 344-355.
GRAY, P. H. H., (1961). Forms and arrangements of scales in species
of Colias ( Lepidoptera : Pieridae). Journal of the New York Entomo-
logical Society, 69( 4) :201-202.
KINDER, E. and F. StiFFERT, (1943). tiber den Feinbau schillernder
Schmetterlingsschuppen vom Morpho-Typ. Biol. Zentr., 63:268.
PORTIER, P., (1932). Sur la structure des ailes des Parnassiens (Lepidop-
teres, Rhopaloceres ) . Comptes Rendus de la Societe de Biologie, 110
(21):465-467.
RICHARDS, O. G., (1944). Notes and news in entomology (Stereoscopic
electron micrographs of Morpho cypris iridescent scales). Entomol.
News, 55(7):190-193.
WATT, W. B., (1964). Pteridine components of wing pigmentation in the
butterfly Colias eurytheme. Nature, 201(4926) : 1326-1327.
YAGI, N. (1954). Note of electron microscope research on pterin pigment
in the scales of pierid butterflies. Annotations Zoologicae Japonenses,
27(3):113-114.
16
HOVANITZ
Fig, 1. — Scene on north side of Sugarloaf Mt., Arabis holboelli var. pine-
torum in foreground.
8(1):16-17, 1969(1970) HABITAT — EUCHLOE
17
Fig. 2. — Closeup of food plant shown in Fig. 1.
HABITAT — Euchloe hy antis andrewsi
Paul A. Opler has recently indicated the distribution of
nearctic Euchloe (J. Res. Lepid. 7(2):65-86). On his map (Fig.
4 as cited), the populations from the San Bernardino mountains,
California, are designated by the name E. hy antis andrewsi.
Collections of this race in July, 1970 were made by the author
at the east end of the San Bernardino mountains on the north
side of Sugarloaf Peak at about 8000 feet elevation (Fig. 1 and
Fig. 2). Females were noted to be laying eggs on Arahis hol-
boellii Hornem. var. pinetorum (Tides.) Roll, which was quite
abundant in the vicinity. Plant identification was kindly made
by Dr. James Hendrickson.
W. Hovanitz
18
CLARK AND DICKSON
J. Res. Lepid.
Fig. 2. Eurema desjardinsi. 1. Imago; 2. Egg; 3. Larva on hatchings; 4. 7th Segment,
1st instar; 5. Head, 1st instar; 6. Larva, 3rd instar; 7. 7th Segment, 2nd instar;
8. Larva, 4th instar; 9. 7th Segment, 4th instar; 10. Larva, final instar; 11. 7th Segment
final instar; 12. Spiracle enlarged; 13. Head, final instar; 14. Pupa; 15. Cremastral
hooks much enlarged; 16. 7th Segment, 3rd instar; Food Plant: Cassia mimosides.
Fig. 1. Eurema hecabe. 1. Imago; 2, Egg; 3. Larva, 1st instar; 4. 7th Segment, 1st instar;
5. Head, 1st instar; 6. Larva, 2nd instar;. 7. 7th Segment, 2nd instar; 8. Larva, 3rd
instar; 9. 7th Segment, 3rd instar; 10. Larva 4th instar; 11. 7th Segment, 4th instar;
12. Larva final instar; 13. 7th Segment, final instar; 14. Spiracle; 15. Head final
instar; 16. Anal comb; 17. Setae, much enlarged; 18. Pupa, cremastral hooks much
enlarged; Food Plant: Cassia mimosides.
Reprint of Fig. 1 and Fig. 2 from
Clark, G. C. and C. G. C. Dickson. 1965.
J. Res. Lepid. 4(4) : 252-257.
8(1):18^19, 1969(1970) SOUTH AFRICAN EUREMA
19
Journal of Research on the Lepidoptera
8(1):20, 1969(1970)
HOVANITZ
HABITAT — ARGYNNIS NOKOMIS
Argynnis nokomis is found throughout the basin and range
country from the eastern side of the Sierra Nevada to the Rocky
Mountains in widely separated isolated spots where there are
cold water seepages, and acid bogs, in the midst of otherwise
alkaline country. As with other Argynnis around the world, the
food plant is probably Viola, though it has not been identified at
this locality.
W. Hovanitz
Fig. 1. — Bog in Round Valley, Inyo Co., California, looking south toward
the Sierra Nevada. June, 1970. Adults fly primarily in late July and August.
20
Journal of Research on the Lepidoptera
8(l):21-36, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1968
BUTTERFLY LARVAL FOODPLANT RECORDS
AND A PROCEDURE FOR REPORTING
FOODPLANTS'
OAKLEY SHIELDS
Department of Entomology, University of California, Davis, Calif ornia
JOHN F. EMMEL
University of California School of Medicine, San Francisco, California
DENNIS E. BREEDLOVE
Department of Botany, California Academy of Sciences,
San Francisco, California
INTRODUCTION
In recent years there has been much interest in the relation-
ships between plants and butterflies (e.g., Brower and Brower,
1964; Ehrlich and Raven, 1965). In much of the past work the
method of recording this data has been inaccurate and unsys-
tematic. The importance of accurately determined larval food-
plants of butterflies has been recognized by some workers but
neglected by many others. Progress in this field has been slow;
as late as 1947 there were a large number of North American
species for which not a single foodplant was known, including
certain common species (Remington, 1947b).
Larval foodplants aid in constructing the biology of the butter-
fly since spatial and temporal distribution, abundance, and some-
times the color pattern of the adult are directly dependent on
foodplants. Thus, one of the keys to the biology of the butterfly
ultimately depends on the precise identification of its larval
foodplant(s). Although some species such as Vanessa cardui
(Linnaeus) and Strymon melinus Hubner utilize a wide variety
of plants, most species appear to be restricted to a few or even
a single plant species. Butterfly foodplants may even help to
determine plant distribution; e.g., Speyeria indicate that Viola is
Hhe term ‘‘foodplant” is used throughout this paper sinee it refers to a
plant that the inseet habitually feeds on, as opposed to “hostplant” which
refers to a plant that the insect lives on ( Torre-Bueno, 1937).
21
22
SHIELDS, ET AL
J. Res. Lepid.
present, a plant that is sometimes not noticeable in a locality at
certain times of the year.
In view of numerous errors now present in the literature, it is
critical that a standardized procedure be established to more
accurately determine foodplant-butterfly relationships, since no
such procedure exists. In this work we propose a procedure for
systematically identifying and reporting plant-butterfly records
so that they can be referred to accurately and with assurance.
We will also discuss past good and bad practices, methods used
to find foodplants, and will report foodplant records for 14
butterfly species based on one season of observation. Additional
records will be reported in future papers.
REVIEW OF LITERATURE ON FOODPLANTS;
VALUE OF FOODPLANTS
Some literature concerning insect-foodplant relationships is
available. A series of papers deal with food selection in phy-
tophagous insects (Brues, 1924; Dethier, 1953, 1954, 1968;
Fraenkel, 1953; Thorsteinson, 1960; Cartier, 1968; Schoonhoven,
1968). Discussions of the effects of available food in relation to
oviposition and larval dispersal (Dethier, 1959a, b), visual and
chemical stimuli used during oviposition (Use, 1937; Cripps,
1947; Fox, 1966; Schoonhoven, 1968), and the variation in selec-
tivity of foodplants (Forbes, 1958; Straatman, 1962a; Stride and
Strattman, 1962) are available for butterflies. Hovanitz and
Chang (1962a, b, 1963a, b, c, d, e, 1964) performed a series of
laboratory experiments with Pieris species, principally Pieris
rapae L., to determine oviposition preferences and responses,
factors affecting foodplant preferences of the larvae, and the
effect of the foodplant onthe larva’s survival and growth rate.
Some work has been done with foodplant specificity in sibling
species of butterflies (Remington and Pease, 1955; Remington,
1958. Emmel and Emmel (1962) discuss factors that limit but-
terfly species to particular foodplants and thus influence the
amount of plant utilization. Downey (1962) found that food-
plant association in Plebejus icarioides ( Boisduval ) may depend
on other factors besides the particular lupine species, such as
pilosity and hybridization in the plant, ant symbiosis, parasites,
competitors, and soil types.
Three major sources to locate butterfly foodplants for North
America are Edwards (1889), Davenport and Dethier (1937),
and Dethier ( 1946 ) . These cite the literature but do not critically
evaluate the foodplants given. J. A. Comstock has published a
series of life history studies of North American butterflies that
8(l):21-36, 1969(1970) FOODPLANT RECORDS
23
includes foodplants, in the Bulletin of the Southern California
Academy of Sciences. Kendall (1959, 1964, 1965, 1966) recorded
foodplants for certain Texas butterflies, and Remington (1952)
reported foodplants for some Colorado species. Detailed work
has been done with the foodplants of one species, Plebejus
icarioides (Downey and Fuller, 1961; Downey, 1962). Work on
foodplants of butterflies in other countries (e.g., Scudder, 1874;
Platt, 1921; Stokoe, 1944; Allan, 1949; Iwase, 1954, 1964; Dick-
son, 1965) may assist in finding new foodplants for North
American species.
Although our knowledge of butterfly foodplants is at a far less
complete state than butterfly taxonomy, foodplants have already
proved to be a valuable tool in interpreting certain evolutionary
trends. Effects of competition and predation on foodplant selec-
tion in butterffies are discussed by Brower ( 1958a, b ) . By analy-
sis of foodplants of three closely related species of Papilio,
Brower (1958b) suggests that competition among the larvae
probably produced restricted and mutually exclusive diets.
(However, D. V. McCorkle, personal communication, found
larvae of two of these, namely Papilio eurymedon Lucas and P.
multicaudata Kirby, feeding on the same Prunus species in
Washington.) Brower (1958a) also found evidence that food
preferences of butterflies that are procryptic and palatable to
birds result from selective pressure favoring those on mutually
exclusive plants due to birds concentrating on a common prey
image. Brower and Brower ( 1964 ) found a strong correlation
between adult butterflies being unpalatable to vertebrate pre-
dators and a narrow range of larval foodplants containing poison-
ous substances. Dethier ( 1941 ) examined various species of
citrus and parsley families and found that these plants have
certain essential oils in common that probably account for their
attractiveness to Papilio larvae. Similarly, Ehrlich nd Raven
(1965, 1967) concluded from a systematic evaluation of plants
eaten by certain butterfly subgroups that butterflies may feed
on plants distantly related phylogenetically but which contain
similar secondary plant substances. From this they suggest that
butterflies and plants are co-evolving. Breedlove and Ehrlich
(1968) found that the seed set of Lupinus amplus Greene was
strikingly reduced by larval infestation of Glaucopsyche lyg-
damus (Doubleday) in one lupine population in Colorado,
indicating that this butterfly could be a strong selective agent on
this plant species. Hovanitz (1949: 351, 353) points out that
man can accelerate the rate of hybridization between two Colias
24
SHIELDS, ET AL
J. Res. Lepid.
species by disturbing the habitat and enabling weeds to encroach.
C alias Christina Edwards thus entered the dwarf willow habitat
of C. gigantea Strecker in southern Canada and C. philodice
Godart entered the Vacciniiim habitat of C. interior Scudder in
northern Michigan following their foodplant invasion along
roadsides.
One practical aspect of knowing the foodplants for butterflies
is in plotting the butterfly’s distribution. For example, Speyeria
nokomis (Edwards), a species usually found in isolated colonies,
can be discovered in new localities by locating herbarium records
for Viola nephrophijlla Greene within its known range and ele-
vation limits.
ERRORS
Past work dealing with butterfly foodplants has often been
imprecise, inadequate, and erroneous. Burns (1964:18), in ascer-
taining Enjnnis foodplants, said ‘‘rampant misidentification is a
serious source of error, hard to detect,” and lightly dismissed
many old records. Downey (1962) said that “considerable error”
exists for butterfly foodplants in the literature. He attributes this
to ( 1 ) data based on single observations and ( 2 ) casual identifi-
cation of the suspected plants. Ehrlich and Raven (1965), in
summarizing foodplant relationships in butterflies, say that
“extreme care has been taken in associating insects with partic-
ular food plants, as the literature is replete with errors and
unverified records.” They mention that despite erratic oviposition
behavior often displayed by butterflies, oviposition records are
frequently considered as foodplant records. Brower (1958b)
pointed out sources of error from evaluating foodplants of three
western United States Papilio species: (1) authors often failed
to indicate whether or not they reared adults from larvae for
positive identification, (2) worn females of the three species
look alike in flight so that oviposition records without capturing
the females are subject to error, and (3) later authors often
quote earlier authors who were mistaken in their information.
Examples of the kinds of errors that are made may help focus
attention on the pitiful state of our knowledge of butterfly food-
plants and may suggest ways to remedy the situation. Tietz
( 1952 ) states that “every effort has been made to list all food-
plants where they are known,” but usually gives no references to
the foodplants listed. He noted Battus philenor ( Linnaeus )
ovipositing on Polygonum scandens L. and thus listed it as a
foodplant. Also, among Euphydryas phaetons (Drury) food
8(l):2l-36, 1969(1970) FOODPLANT RECORDS
25
plants were listed Ribes, Corylus, and Fraxinus, all unlikely to
serve as foodplants. Garth and Tilden (1963) did not designate
foodplant species because it “would have prolonged the list
unduly” and list genera for the most part. Edwards (1868-1872)
reported Polijgonia zephijrus (Edwards) on Azalea occidentalis
(now known as Rhododendron occidentale (T. & G.) Gray) and
later (1884) corrected his mistake in two places, saying that the
larva and pupa that were drawn referred to Polygonia faunus
rusticus (Edwards). Despite this correction many texts since
have continued to list Azalea as a zephyrus foodplant. One of
the present authors (JFE) reported (1962) that Lycaena cupreus
(Edwards) larvae were found on Calyptridium umbellatum
( Torr. ) Greene; they were not reared to adult. Despite the fact
that a female cupreus was seen to oviposit on the Calyptridium
earlier in the season, it is probably not the foodplant; a later
investigation of the area in 1965 by JFE revealed that a Rumex
species, probably the true foodplant, was growing abundantly
among the Calyptridium. The larvae that were found are now
thought to have been Strymon melinus, but this is only specula-
tion. This example emphasizes the need to follow through on
observations of oviposition before considering a plant as a food
source. One wonders how certain peculiar errors ever developed
in the first place, such as Neophasia terlooti Behr feeding on
“mistletoe” (Forbes, 1958). Stokow (1944) and Allan (1949)
did not distinguish between laboratory and field rearings for
species of foodplants of British butterflies.
Species are often said to feed on a common plant, implying
that a particular species is a general feeder on that group of
plants. For example, references to Polygonia satyrus (Edwards)
on “nettle”, Satiyrium sylvinus (Boisduval) on “willows”, and
many satyrines and hesperiids on “grasses” are common. The
inaccuracy of such statements is pointed out by the fact that not
one specific grass genus, let alone species, is known for a North
American satyrine. (However, N. McFarland, in litt., reports a
Cercyonis larva on Dactylis glomerata L. 5 miles W. of Gorvallis,
Oregon.) A sedge may be the foodplant of the satyrine Eupty-
chia mitchellii (French) (McAlpine, Hubbell, and Pliske, 1960)
and sedges are strongly suspected for at least one species of
Oeneis (JFE and OS, personal observation).
Brower ( 1958b ) traced one error down. Gomstock had re-
ported the foodplant for Papilio rutulus Lucas as “hop”, which
was reported elsewhere as Humulus when he meant Ptelea
Baldwinii T. & G. (Hop-Tree). In Philotes, the Eriogonum food-
26
SHIELDS, ET AL
/. Res. Lepid.
plant is quite specific for any given population, yet Downey in
Ehrlich and Ehrlich ( 1961 ) states that they feed on '"Eriogonurri’ .
One problem with erroneous foodplant determinations is that
it is difficult to improve them or even sometimes to distinguish
them from legitimate records when no documentation accom-
panies the statement. Sometimes apparently legitimate records
by reputable workers are erroneous, such as W. H. Edwards re-
porting Papilio inclra Reakirt as feeding on Artemisia dracuncu-
Ills L. ( Emmel and Emmel, 1963 ) . It will be a long, slow process
to weed out erroneous records, and it would be advisable to
duplicate legitimate records with adult and plant reference
material. Records suspected to be erroneous should be corrected
when new data dictates it. For example, Davenport and Dethier
( 1937 ) gave Lotus glaber Greene and Astragalus sp. as well as
Purshia glandulosa Curran reported in the literature as food-
plants for Callipsyche hehrii (Edwards). The reference to
Purshia is well documented (Comstock, 1927, 1928). The range
of the adult corresponds to that of Purshia and the lars^ae have
subsequently been found on Purshia but the other two records
have never been duplicated. A look at the original source
(Williams, 1908) reveals that the Lotus and Astragalus records
refer to '‘Lijcaena hehrii \ plainly a species of “blue” from the
context.
At a somewhat lower level, subspecies of plants are not often
given, although such a reference can be important. For example,
Papilio indra fordi Comstock & Martin was originally described
as feeding on Cijmopterus panamintensis Coult. & Rose, although
it does not occur on the nomotypical subspecies but rather only
on the subspecies acutifoliiis (Coult. & Rose) Munz (JFE, un-
published). Sometimes certain records are common knowledge
yet are not published; this is also a type of error.
Some authors are of the opinion that choice of foodplants is an
indication of butterfly relationships (Ae, 1958; Forbes, 1958;
Garth and Tilden, 1963:16). Garth and Tilden (1963) cite as an
example certain Colias species that feed on Vaccinium instead of
“preferred” legumes and therefore should be set apart from
others of their genus. However, there is some evidence that this
is a conditional argument. For example, considering morphologi-
cal characters, Papilio indra and its subspecies, strictly Umbelli-
ferae feeders, are not closely related to the P. machaon L. species
complex which has species that feed on Umbelliferae, Compos-
itae {Artemisia dracunculus for P. hairdii Edwards), and Ruta-
ceae {Thamnosma montana Torr. & Frem. for P. rudkini Com-
8(l):21-36, 1969(1970) FOODPLANT RECORDS
27
stock). Using foodplants here for taxonomic purposes, that would
make P. indr a closer to the P. machaon complex than either P.
hair da or P. rudkini is. The potential of foodplant relations as
data for butterfly classification is discussed by Downey (1962).
REPORTING PROCEDURE AND COLLECTING METHODS
To help overcome the mistakes made in the past in reporting
foodplants, we wish to establish certain guide-lines to follow.
Several such attempts have been made in the past. Remington
(1947a) proposed that the Lepidopterists’ Society would have a
botanist available to determine foodplants; however, the idea
apparently did not materialize. Opler ( 1967 ) , in giving new
foodplants for Anthocaris sara Lucas and A. lanceolata Lucas,
confirmed the determinations with a botanist, gave exact locality
and date that the plant was collected, gave the circumstances
under which the plant was found to be a food source, and even
reported the determination down to “varieties” (= subspecies).
However, no place of deposition was assigned for the plants or
immatures. Remington (1952) deposited foodplants at a desig-
nated herbarium.
Foodplants should be determined by a competent botanist
and placed on file with a recognized herbarium specifically re-
ferred to for later inspection if ever needed. (Herbaria of the
world are listed in Lanjouw and Stafleu, 1959, with their proper
abbreviations). Some groups of plants must be determined by
a specialist. Herbarium records are always mandatory. Certain
groups such as Agave and Lupinus as yet have not been revised
satisfactorily. We hope that eventually all North American but-
terflies will have their foodplants on file in herbaria for future
reference.
A plant press should be part of the standard equipment of the
lepidopterist concerned with butterfly biology. Flowers and/or
fruit are essential for determination of most plant species. In
instances where oviposition or immatures occur on plants with
no flowers or fruit, leaf characteristics should be carefully com-
pared with surrounding plants (to be used for specimens), and
this should be stated when recording the plant. When a female
oviposits on a plant species that is not in bloom, it is sometimes
necessary to return to the exact spot later in the season or the
following year to collect the same plant with flowers or fruit
(the plant should be marked). Also, plants that ovipositing
females are “interested in” may also be the clue to finding the
28
SHIELDS, ET AL
/. Res. Lepid.
foodplant; suspected foodplants, properly documented, are val-
uable to report since they assist in finding new foodplants.
Just as preserving foodplants is a necessity, preservation of
the butterfly stage connected with the foodplant is extremely im-
portant. Whether it was an ovipositing female or adults ulti-
mately reared from in situ larvae, or^ eggs, larvae, or pupae
compared with known species, the material should be preserved
and deposited in a designated museum for later reference by
future workers. This is particularly important in case of future
revisions and the naming of new subspecies.
Giving the locality of the foodplant is important because
different foodplants are often used in different localities, and
the same species that serves as a foodplant in one locality may
not serv^e as a foodplant in another locality (Downey, 1962).
Vegetation type is important to report. For example, we found
Satyrium fuliginosum (Edwards) only in sagebrush areas even
though its foodplant, a Liipiniis species, occurred in other habi-
tats. The condition of the foodplant is often important. Fre-
quently species will prefer to oviposit on seedlings of the food-
plant or on plants without flowers. Vanessa virginiensis (Drury)
oviposits on Gnaphalium seedlings (Dethier, 1959a) and Vanes-
sa cardui will oviposit on small, second growth thistles (Keji,
1951).
Evidence of feeding may be important in determining new
localities for a species when no immatures or adults are present.
For example, Megathyminae larvae construct “trap doors” and
“tents”, and Papilio bairdii larvae strip Artemisia dracunculus
stems of leaves and deposit a characteristic type of feces on the
ground.
Surprisingly little has been written about methods of locating
foodplants of butterflies. Kuzuya (1959) told how to locate
theclini eggs in winter in Japan, which helps to locate their
foodplants. McFarland (1964) discussed methods of collecting
Macrolepidoptera larvae. In the future, it would be helpful to
know the location of eggs on the foodplant and what part of the
plant the larvae eat, to assist in finding immatures and food-
plants. For example, we found Lycaena eggs in stem axils and
Euphydryas egg masses only on the underside of the leaves.
Larvae may feed on certain parts exclusively such as young
leaves, flowers, or bark. Also, the manner in which the eggs are
laid is important (singly, clusters, or small groups).
The behavior of females is often a clue in discovering food-
plants. A female repeatedly alighting on the same plant species
8(l):21-36, 1969(1970) FOODPLANT RECORDS
29
and curling her abdomen toward the plant should be watched.
If the female does not lay eggs on the plant, the plant should be
checked anyway for eggs from other females. Certain females
such as Speyeria, Parnassius, and Satyrium fuliginosum do not
always oviposit directly on the foodplant, so that choice of food
with these is the responsibility of the young larva. Hesperia
Undseyi Holland oviposits on lichens or some other substrate;
the larvae must select the proper grass species (MacNeill,
1964:32). Female oviposition on a plant may not necessarily
mean the plant is a foodplant. Examples of “mistakes” by fe-
males are well known. Coolidge (1925) found Hylephila phy-
laeus (Drury) ovipositing on grasses, rocks, twigs, and even a
paved street. Speyeria oviposit on dried leaves (Ritchie, 1944),
various plants (Guppy, 1953), and Artemisia bark (Durden,
1965), but the larvae eat leaves of Viola species. There are ex-
amples of butterfly species ovipositing on introduced plants on
which the resultant larvae do not survive (Remington, 1952;
Brower, 1958b; Brooks, 1962; Straatman, 1962b; Sevastopulo,
1964).
In the genera Euphydryas, Chlosyne, and Phyciodes, it is
sometimes easier to search for larval webs on suspected food-
plants in summer or fall after the adults have flown than it is to
follow females or to search for eggs. Newcomer (1967) found
larvae of Chlosyne hoffmanni manchada Bauer on Aster con-
spicuus Lindley by looking for larval webs in July after the
adults had flown.
Knowing only one species’ foodplant can be useful in locating
foodplants for other members of the same genus (e.g., Speyeria
and Euphydryas). Sometimes it may be helpful to locate areas
where few possible foodplants are available so that the foodplant
can be located easily. For example, Ochlodes yuma (Edwards)
flies in some areas where its foodplant, Phragmites communis
Trin., is the only grass present.
In problem groups such as Satyrinae, it may be necessary to
place possible foodplants with caged females for clues or to
statistically analyze the numbers of young larvae that crawl
toward, feed on, and remain on a variety of plant species placed
in a petri dish.
Often the areas where females oviposit are away from the
flight areas of the males; locating such areas of female concen-
tration increases the probability of finding foodplants. For ex-
ample, we found an area where Colias scudderii Reakirt females
30
SHIELDS, ET AL
J. Res. Lepid.
were ovipositing on low-growing Salix plants in only one small
section of a willow bog in Colorado.
Knowing when is the best time to find foodplants can be
useful. Langston (1963) states that the appearance of Eriog-
onum-f ceding Philotes adults is correlated with the early full-
bloom of Eriogonum. Thus a knowledge of the blooming time in
this case helps to locate immatures and their foodplants.
Using a technique suggested by Mr. Christopher Henne
( personal communication ) , we have had success in finding
lycaenid larvae in flowerheads by drying out picked fiowers of
the suspected plant, thus forcing the larvae to crawl up the sides
of the container in search of fresh food.
DEPOSITIONS AND DETERMINATIONS
Foodplant records have been recorded intermittently by two
of us ( JFE and OS ) since 1967. The number by the plant is the
collector’s number (for J. F. Emmel) for the plant. The de-
posited butterfly material is labelled to include this number.
The herbarium sheets with the exception of the Umbelliferae
will be deposited with their respective species at the Dudley
Herbarium, Stanford University, Stanford, California (DS); the
Umbelliferae will be deposited at the U. C. Berkeley Herbarium,
Berkeley, California (UC); and the preserved butterfly material
will be deposited at the Los Angeles County Museum, Los An-
geles, California.
Most of the plants were identified by one of us (DEB).
Species of the genus Eriogonum were identified by Dr. James
L. Reveal, Department of Botany, University of Maryland,
College Park, Maryland, and the Umbelliferae were determined
by Dr. Lincoln Constance, Department of Botany, University of
California, Berkeley, California.
We wish to thank Mr. Noel McFarland for his helpful criti-
cisms of the manuscript. This work was supported by a grant
from the Allyn Foundation, Chicago, Illinois, for the summers of
1968-1969, and N.S.F. Crant no. GB-5645, for the summer of
1967.
FOODPLANT RECORDS
(All collected by JFE and OS unless otherwise noted. Plant
genera of the world can be placed to family by reference to
Willis, 1966. )
PIERIDAE
I. Colias alexandra Edwards. Wasatch Plateau, 10,000', near
Mt. Sanpete, E. of Ephraim, Sanpete Co., Utah, 31 July 1967,
8(l):21-36, 1969(1970) FOODPLANT RECORDS
31
female oviposited at 11:30 AM MST on leaf of Astragalus
miser Dougl. ex Hook. (Leguminosae), /. F. Emmel 25 (DS).
2. Colias meadii Edwards. Cottonwood Pass, 12,200', Chaffee
Co., Colo., 28 July 1967, female oviposited between 8:20-
9:30 AM MST on leaf underside of Trifolium dasyphyllum
Torr. & Gray ( Leguminosae ) , /. F. Emmel 22 ( DS ) .
3. Euchloe ausonides coloradensis (H. Edwards). (A) Dry
meadow at 9600', Gothic, Gunnison Co., Colo., 10 July 1967,
female oviposited at 10:00 AM MST on flower bud of Arabis
drummondi Gray ( Cruciferae ) , /. F. Emmel 6 (DS). (B)
North side of Schofield Pass, 10,400', Gunnison Co., Colo.,
14 July 1967, female oviposited at 2:00 PM MST on flower
bud of Descurainia calif ornica (Gray) O. E, Schulz (Cruci-
ferae), /. F. Emmel 11 (DS). (C) Schofield Pass, 10,500',
Gunnison, Co., Colo., 18 July 1967, female oviposited at 1:30
PM MST on flower bud of Descurainia calif ornica (Gray)
O. E. Schulz (Cruciferae), /. F. Emmel 13 (DS).
4. Pieris napi (Linnaeus). (A) East River at 9600', in wet
meadow among willows, near Gothic, Gunnison Go., Colo.,
10 July 1967, female oviposited at 10:00 AM MST on leaf
underside of Cardamine cordifolia A. Gray (Cruciferae),
/. F. Emmel 7 (DS). (B) Meadow 54 mile S. Brush Creek
Cow Camp, 9000' near the East River, Gunnison Co., Colo.,
12 July 1967, female oviposited at 10:00 AM MST on leaf
underside of Thlaspi arvense L. (Cruciferae), /. F. Emmel 10
(DS). (C) Cement Creek, Gunnison Co., Colo., 18 July
1967, female oviposited at 2:00 PM MST on leaf underside
of Thlaspi arvense L. (Cruciferae), /. F. Emmel 14 (DS).
5. Pieris occidentalis Reakirt. (A) East slope of Belleview
Mountain, 11,700', near Schofield Pass, Gunnison Co., Colo.,
25 July 1967, female oviposited at 11:30 AM MST on leaf
underside of Thlaspi alpestre L. (Cruciferae), /. F. Emmel 21
(DS). (B) Rockslide above Island Lake, 10,500', Ruby Mts.,
Elko Co., Nev., 8 Aug. 1967 (collectors JFE, OS, and S. Ellis),
female oviposited at 10:15 AM PST on leaf underside of
Draha cuneifolia Nutt .ex T. &. G. (Cruciferae), /. F. Emmel
32 (DS).
NYMPHALIDAE
1. Chlosyne acastus Edwards. In washes along road, 9 miles W.
of Vernal on U.S. Hwy. 40, Uintah Co., Utah, 21 Aug. 1967
(collectors JFE, OS, and S. Ellis), two larvae on plant stems,
pair reared to adult (emerged 22 Feb. 1968, male; 21 Feb.
1968. female), on Machaer anther a viscosa (Nutt.) Greene
32
SHIELDS, ET AL
J. Res. Lepid.
( Compositae ),J.F. Emmel 39 ( DS ) .
2. Chlosyne palla calydon Strecker, On grassy slope with aspen,
sagebrush, and Castilleja, near Brush Creek Cow Camp above
the East River, 9100', Gunnison Co., Colo., 27 Aug. 1967,
larva in web near base of stems (adult formed inside pupa,
a male; genitalia identical to C. palla from Colorado in the
Los Angeles County Museum and to the drawing in Ehrlich
and Ehrlich, 1961 ) , on Erigeron speciosus ( Lindl. ) DC
( Compositae ), J. F. Emmel 41 ( DS ) .
3. Polygonia zephyrus (Edwards) Charleston Park, 8300',
Charleston Mts., Clark Co., Nev., 31 Aug. 1967, larva on
stem (male emerged 16 Sept. 1967) of Ribes cereum Dougl.
( Saxifragaceae ) , /. F. Emmel 45 (DS).
4. Speyeria atlantis clodgei (Gunder). Lost Prairie, W. of
Santiam Pass on U.S. Hwy. 20, Linn Co., Ore., 12 Aug. 1967
(collectors JFE, OS, and S. Ellis), female oviposited on leaf
underside (female reared from this female, emerged 6 Apr.
1968) of Viola bellidifolia Greene (Violaceae), /. F. Emmel
36 (DS).
LYCAENIDAE
1. Glaucopsyche lygdamus oro Scudder. Large, open, dry mea-
dow, north side of Schofield Pass, 10,400', Gunnison Co.,
Colo., 14 July 1967, female oviposited at 1:45 PM MST on
flower bud of Lupinus ammophilus Greene (Leguminosae),
/. F. Emmel 12 (DS).
2. Plebejus argyrognomen ricei (Cross). (A) Lost Prairie, W.
of Santiam Pass, on U.S. Hwy. 20, Lifm Co., Ore., 12 Aug.
1967 (collectors JFE, OS, and S. Ellis), female oviposited at
12:15 PM PST on stem near base of plant of Vicia exigua
Nutt. (Leguminosae), /. F. Emmel 38 (DS). (B) Lost
Prairie, W. of Santiam Pass, on U.S. Hwy. 20, Linn Co., Ore.,
12 Aug. 1967 (collectors JFE, OS and S. Ellis), female
oviposited at 12:30 PM PST on stem near base of plant of
Lathyrus torreyi Gray ( Leguminosae ) , /. F. Emmel 37 ( DS ) .
3. Plebejus saepiolus ( Boisduval ) . ( A ) Crested Butte Cemetery,
8900', Crested Butte, Gunnison Co., Colo., 12 July 1967,
female oviposted inside flower-head between flowers of
Trifolium repens L. (Leguminosae), /. F. Emmel 8 (DS).
(B) Crested Butte Cemetery, 8900', Crested Butte, Gunnison
Co., Colo., 12 July 1967, female oviposited inside flower-head
between flowers of Trifolium longipes Nutt. (Leguminosae),
/. F. Emmel 9 (DS). (C) Trail from Pine Creek Camp to
Mt. Jefferson, 10,500', Toquima Range, Nye Co., Nev., 4 Aug.
8(1):21^36, 1969(1970) FOODPLANT RECORDS
33
1967 (collectors JFE and S. Ellis), female oviposited at 1:00
PM PST on side of flower of Trifolium monanthum Gray
( Leguminosae ) , J. F. Emmel 29 ( DS ) .
HESPERIIDAE
1. Hesperia uncas Edwards. Hilltop 2 miles S. of Gunnison,
8000', Gunnison Co., Colo., 27 Aug. 1967, female oviposited
at 11:10 AM MST on leaf underside of Bouteloua gracilis
(HBK. ) Lag. (Gramineae), /. F. Emmel 42 (DS).
2. Thorybes mexicana nevada Scudder. Open dry meadow near
Crested Butte Cemetery, 8900', Crested Butte, Gunnison Co.,
Colo., 30 June 1967, female oviposited at 10:55 AM MST on
leaf underside of Lathyrus leucanthus Rydb. (Leguminosae),
/. F. Emmel 2 (DS).
LITERATURE CITED
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ALLAN, P. B. M., 1949. Larval foodplants, a vade-mecum for the field
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BROOKS, J., 1962. Foodplants of Papilio palamedes in Georgia. /. Lepid.
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BROWER, L P. 1958a. Bird predation and foodplant specificity in closely
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BROWER, L. P., and J. V. Z. BROWER, 1964. Birds, butterflies, and
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BURNS, J. M., 1964. Evolution in skipper butterflies of the genus Erynnis.
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COMSTOCK, J A., 1927. Studies in Pacific Coast Lepidoptera (cont. ),
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1928. Studies in Pacific Coast Lepidoptera (continued). Bull. So.
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COOLIDGE, K. R., 1925. Life history studies of some California Rhopalo-
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1946. Supplement to the bibliography of the described life-histories
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1953. Host plant perception in phytophagous insects. Trans. Ninth
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34
SHIELDS, ET AL
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1954. Evolution of feeding perferences in phytophagous insects.
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1959b. Food-plant distribution and density and larval dispersal as
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DICKSON, C. G. C., 1965, Recently observed food-plants of some South
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15:34-42.
DURDEN, C. J,, 1965. Speyeria callippe and Artemisia, a possible food-
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EDWARDS, H, 1889. Bibliographical catalogue of the described trans-
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EHRLICH, P. R,, and A. H. EHRLICH, 1961. How to know the butter-
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1967. Butterflies and plants, Sci. Amer., 216( 6) : 105-113.
EMMEL, J. F., and T. C. EMMEL, 1963. Larval food-plant records for
six Western Papilios. J. Res. Lepid., 1:191-193.
EMMEL, T. C., and J. F. EMMEL, 1962. Ecological studies of Rhopalo-
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fly associations and distributional factors. J. Lepid. Soc., 16:23-44.
FORBES, W. T. M., 1958. Caterpillars as botanists. Proc. Tenth Inter.
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FOX, R. M., 1966. Forelegs of butterflies. I. Introduction: chemoreception.
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FRAENKEL, G., 1953. Insecets and plant biochemistry. The specificity of
foodplants for insects. Proc. Fourteenth Inter. Congr. Zoo/., 1:383-387.
GARTH, J. S., and J. W. TILDEN, 1963. Yosemite butterflies. J.
Res. Lepid., 2:1-96.
GUPPY, R., 1953. Rearing Speyeria in captivity. Lepid. News, 7:56.
HOVANITZ, W., 1949. Increased variability in populations following na-
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Simpson (ed.) Genetics, paleontology, and evolution, Princeton Uni-
verity Press, Princeton. 474 p.
HOVANITZ, W., and V. C. S. CHANG, 1962a. The effect of various food
plants on survival and growth rate of Pieris. J. Res. Lepid., 1:21-42.
1962b. Three factors affecting larval choice of ood plants. /. Res.
Lepid., 1:51-61.
1963a. Change of food plant preference by larvae of Pieris rapae
controlled by stain selection, and the inheritance of this trait. J. Res.
Lepid., 1:163-168.
1963b. Comparasion of the selective effect of two mustard oils
and their glucosides to Pieris larvae, J. Res. Lepid., 2:281-288.
1963c. The effect of hybridization of host-plant starins on growth
rate and mortality of Pieris rapae. J. Res. Lepid., 1:157-162.
8(l):21-36, 1969(1970) FOODPLANT RECORDS
35
1963d. Ovipositional preference tests with Pieris. J. Res. LepicL,
2:185-200.
1963e. Selection of allyl isothiocyanate by larvae of Pieris rapae
and the inheritance of this trait. J. Res. Lepid., 1:169-182.
1964. Adult ovipositional responses in Pieris rapae. J. Res. Lepid.,
3' 159-172
HOVANITZ, W., V. C. S. CHANG, and G. HONCH, 1963. The effective-
ness of different isothiocyanates on attracting larvae of Pieris rapae.
J. Res. Lepid., 1:249-259.
ILSE, D., 1937. New observations on responses in colours in egg-laying
butterflies. Nature (London), 140:544-545.
IWASE, T., 1954. Synopsis of the known life-histories of Japanese butter-
flies. Lepid. News, 8:95-100.
1964. Recent foodplant records of the Loochooan butterflies. J. Lepid.
Soc., 18:105-109.
KEJI, J. A., 1951. Oviposition observations. Lepid. News, 5:69.
KENDALL, R. O, 1959 More larval foodplants from Texas. J. Lepid. Soc.,
13:221-228.
1964. Larval foodplants of twenty-six species of Rhopalocera (Papi-
lionidae) from Texas. /. Lepid. Soc., 18:129-157.
1965. Larval food plants and distribution notes for twenty-four Texas
Hesperiidae. J. Lepid. Soc., 19:1-33.
1966. Larval food plants for five Texas Hesperiidae. J. Lepid Soc.,
20:35-41.
KUZUYA, T., 1959. The breeding of the Theclini and collecting their eggs
in winter. /. Lepid. Soc., 13:175-181.
LANGSTON, R. L., 1963. Philotes of central coastal California (Lycae-
nidae). /. Lepid. Soc., 17:201-223.
LANJOUW, J., and F. A. STAFLEU, 1959. Index herbariorum. Part I.
The herbaria of the world. 4th ed. Kemink and Zoon; Utrecht, Nether-
lands. 249 p.
McALPINE, W. S., S. P. HUBBELL, and T. E. PLISKE, 1960. The
distribution, habits, and life history of Euptychia mitchellii (Satyridae).
/. Res. Lepid., 14:209-226.
McFarland, N., 1964. Notes on collecting, rearing, and preserving larvae
of Macrolepidoptera. J. Lepid. Soc., 18:201-210.
MacNEILL, C. D., 1964. The skippers of the genus Hesperia in western
North America with special reference to California ( Lepidoptera:
Hesperiidae). Univ. Calif. Pub. Ent., 35:1-230.
NEWCOMER, E. J., 1967. Early stages of Chlosyne hoffmanni manchada
( Nymphalidae). J. Lepid .Soc., 21:71-73.
OPLER, P. A., 1967. New host plant records for Anthocaris (Pieridae).
J. Lepid. Soc., 21:212.
PLATT, E. E., 1921. List of foodplants of some South African lepidopterous
larvae. So. African J. Nat. Hist., 3:65-138.
REMINGTON, C. L., 1947a. Host plant identification. Lepid. News, 1:25.
1947b. Life history studies I. The importance of life history investiga-
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1952. The biology of Nearctic Lepidoptera. I. Foodplants and life-
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REMINGTON, C. L., and R. W. PEASE, JR., 1955. Studies in foodplant
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36
SHIELDS, ET AL
J. Res. Lepid.
SCHOONHOVEN, L. M,, 1968, Chemosensory bases of host plant selection.
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SCUDDER, S. H., 1874. The food-plants of European butterflies. Canad.
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SEVASTOPULO, D. G., 1964. Lepidoptera ovipositing on plants toxic to
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STOKOE, W. J., 1944, The caterpillars of the British butterflies, including
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STRAATMAN, R., 1962a. A hybrid between Papilio aegeus aegeus and
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1962b. Notes on certain Lepidoptera ovipositing an plants which are
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BOOKS:
NOTICES
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastiinil Choc. Paul Hamlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophical
Library, N.Y.
WANTED:
Brephidium exilis, B. fea, B. isophthalma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, Galifornia 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERIGA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systematics, biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
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irtay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR plea.se write notices on a
3x5 card in the form desired and they will be printed in the
next following is.sue of the JOURNAL.
Volume 8
Number 1
March, 1969
IN THIS ISSUE
Scanning electron microscopy
on wing scales of Colias eurytheme.
John M. Kolyer and Anne Marie Reimschuessel 1
Habitat — Euchloe hyantis andrewsi W. Hovanitz 16
South African Eurema
G. C. Clark and C. G. C. Dickson 18
Habitat — Argynnis nokomis W. Hovanitz 20
Butterfly larval foodplant records
and a procedure for reporting foodplants.
O. Shields, J. F. Emmel, and D. E. Breedlove 21
Volume 8 Number 2 June, 1969
THE JOURHAL
©F RESEARCH
©HJ THE LEFI©©RTERA
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Journal of Research on the Lepidoptera
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© Copyright 1969
NOMENCLATURE OF WING VEINS AND CELLS
LEE D. MILLER
Allyn Museum of Entomology
712 Sarasota Bank Bldg.
Sarasota, Florida 33577
Ideally the naming of veins and the cells between them
should be uniform and in at least general agreement; unfor-
tunately, in surveying the literature one soon finds that such is
not the case. There are many systems for naming veins and
almost as many for cell nomenclature. Each author obviously
uses that system most familiar to him, usually disregarding other
schemes, and thus adds to the confusion of the reader. This
bewilderment is most apparent when a non-lepidopterist at-
tempts to use some of the systems that are purely oriented to
Lepidoptera and that bear little or no relationship to the schemes
employed for other orders of insects. I therefore present this
paper in an attempt to unravel some of the confusion generated
by the differences in these diverse systems, but I will also engage
in some “evangelism” in behalf of that scheme that I feel is most
advantageous.
The references below to the different workers employing the
various systems only deal with those works on the New World
butterflies — the reconciliation of the schemes used in papers on
moths will have to be done by another author.
THE HERRICH SCHAFFER SYSTEM
One of the earliest systems for naming veins, and certainly the
oldest one that has survived into the relatively modern literature,
was devised by Herrich-Schaffer over a century ago and first
used in his writings. As shown in Fig. 1, in this system the most
anterior forewing vein is the costa ( C ) , the next five are branches
of the subcosta (from the anterior one, SCi, SCg, SC3, SC4 and
SC5), the following two are the oher-radius (OR) and unter-
radius (UR), the next three are branches of the medius (M3,
37
SMITHSONIAN
INSTITUTION
38
LEE MILLER
J. Res. Lepid.
Fig. 1.— Venation of a hypothetical butterfly according to the Herrich-
Schaffer system. The abbreviations are explained in the text.
No designation for extradiscal cells in the system
8(2):37-48, 1969(1970)
VEINS AND CELLS
39
M2 and Ml, from anterior to posterior; the rationale here being
that Ml arises nearer the base than does M25 etc. ) , and the most
posterior vein that is present in all butterflies is the sub-medius
(SM). If the small vestigial vein posteriad of SM is present, as
it is in the Papilionidae, it is known as the first anal vein (lA).
The hindwing venation is as follows: the short, spur-like vein
near the base and anteriad of the first main vein is the precosta
(pc), the anterior main vein is the costa (C), the second the
subcosta (SC), the next two the oher-radius (OR) and unter-
radius (UR), the following three the branches of the medius
(respectively, Mg, M2 and Mi), the next one (the most anterior
one not connected to the discal cell) the sub-medius (SM) and
the most posteriad vein is the first anal ( 1 A ) . On both wings the
anterior vein delimiting the discal cell is the subcosta and the
posterior vein the medius. The forewing crossveins between SC5
and OR, OR and UR, UR and Mg are the first (upper), second
(middle) and third (lower) discocellulars (respectively, udc,
mdc and ldc).*The hindwing crossveins between SC and OR,
OR and UR are the first or upper (udc) and second or lower
(Idc) discocellulars, respectively.
There are no provisions for naming cells in this system, except
for the discal cell (D) which is the same in all schemes.
Although the Herrich-Schaffer system is not in current usage,
it is of interest to American workers since it is employed in those
sections of Seitz Macrolepidoptera of the World contributed by
Fruhstorfer, Haensch, Rober and in some of Weymer’s discus-
sions. This system is also the one used by Godman and Salvin in
the Biologia Centrali- Americana, and the venation drawings of
Ithomiidae in Holland's Butterfly Book were taken from another
author using the Herrich-Schaffer system.
THE “INDIAN” SYSTEM
The “Indian” system, used chiefly by de Niceville and Moore
in their various writings on the butterflies of the Indian region,
affects Americans only peripherally in comparisons of the Old
and New World faunas. This system differs very little from that
of Herrich-Schaffer: OR and UR of the forewing are designated,
respectively, Discoidal 1 and Discoidal 2; hindwing vein SC of
Herrich-Schaffer is the first subcostal (SCi) in the present sys-
tem, OR is the second subcostal ( SC2 ) , UR is the Discoidal and
lA is the Internal nervule.
40
LEE MILLER
/. Res. Lepid.
Fig. 2. — Venation and extradiscal cells of a hypothetical butterfly according
to the Rothschild-Jordan system. The single symbols represent the names
of veins, and the double ones (i. e., Mi -M2) represent cells, as explained
in the text.
8(2);37-48, 1969(1970)
VEINS AND CELLS
41
THE ROTHSCHILD-JORDAN SYSTEM
The Rothschild- Jordan system (Fig. 2), based in part on the
Herrich-Schaffer system, was the most comprehensive one pro-
posed to its time. In this scheme the most anterior forewing vein
is again the costa (C), the next five branches of the subcosta
(SCi, SC2, SC3, SC4 and SCg, from anterior to posterior), the next
three are branches of the radius (Ri, Rs and R3, from the most
anteriad), the following two branches of the medius (Mi the
anterior and M2 the posterior), and those veins arising posteriad
of the cell are the submedians (SM); the most anterior of these
(SMi) is considered absent and represented by only a fold, the
one present in all Lepidoptera is SM2 and the tiny, posterior spur
is SM3. The forewing crossveins between SC5 and Ri, Ri and
Ro, R2 and R3 are, respectively, the upper (udc), middle (mdc)
and lower (Idc) discocellulars. On the hindwing the short basal
spur anterior to the main veins is the precostal vein (pcv), the
anterior main vein is the costa (C), the second the second sub-
costa (SC2, SCi being considered absent), the next three are
branches of the radius (from the anterior one, Ri, R2 and R3),
the following two are branches of the medius (Mi anteriad and
M2 posteriad) and those veins arising posteriad of the discal cell
are the submedians (SM): the anterior SMi is only a fold in
most butterflies, whereas the middle SM2 and the posterior SM3
are always present. The upper (udc), middle (mdc) and lower
(Idc) disocellulars delimit and end of the discal cell between
veins SC2 and Ri, Ri and R2 and R2 and R3, respectively
The naming of the spaces between the veins outside the discal
cell (D) was formalized in the Rothschild- Jordan system with
great precision. The cells are named for the veins bounding
them, so that the space between veins Mi and M2 is denoted
M1-M2, for example. The cells anteriad of C and posteriad of
the last SM are, respectively, the costal cell and the SM cell.
The Rothschild- Jordan system was used widely by authors
in the last decade of the last century and the first thirty years
of this one. In works pertaining to the American butterflies
Holland adopted the system in The Butterfly Book (with the
exception of the venation drawings of ithomiids mentioned in
the discussion of the Herrich-Schaffer system), and the Roth-
schild-Jordan scheme is employed in those parts of Seitz authored
42
LEE MILLER
J. Res. Lepid.
Fig. 3. — Venation and extradiscal cells of a hypothetical butterfly according
to the “English”, or numerical, system. Those symbols preceded by “Int”
refer to the cells, and the numbers alone refer to the veins, as explained
in the text.
8(2);37-48, 1969(1970)
VEINS AND CELLS
43
by Seitz, Jordan and Draudt. Naturally, the system is employed
in Rothschild and Jordans revision of the American Papilio.
THE ^‘ENGLISH”, OR NUMERICAL, SYSTEM
The “English”, or numerical, system (Fig. 3) is a totally
artificial system whose major advantage is its great simplicity.
In this scheme the main veins of both wings are named from the
most posterior to the most anterior: hence, all butterflies have
veins 1-12 on the forewing and veins 1-8 on the hindwing. The
only source of confusion concerning the nomenclature of veins
is in the designations of those which arise posterior to the discal
cells (D) of both wings. On the forewing the vestigial vein
posteriad of vein 1 is denoted as la (the fold between veins
1 and 2 represents the primitive vein lb), and on the hindwing
the possible veins posteriad of the cell are veins Ic, 1 and la,
from the cell to the inner margin. The hindwing precostal vein
(pvc) of other systems bears no special designation in the
“English” system and the discocellular veins ( udc, mdc and Idc )
of both wings are as in the Rothschild-Jordan system.
The naming of the extradiscal spaces (Int. ) is equally simple:
the cells are named for the veins posteriad of them — thus the
cell between veins 6 and 7 is known as Int. 6. The only apparent
inconsistency concerns the spaces on either side of vein 1 of
both wings. The cell anteriad of forewing vein 1 is Int. lb, and
the one posterior to vein 1 is Int. la. The hindwing cells from
vein 2 and the inner margin are Int. Ic, Int 1 and b respect-
ively.
The numerical system is followed by most British and some
American writers. It is chiefly of interest to workers on American
butterflies because of its use by Weymer in Seitz (but not in his
discussions where he uses the Herrich-Schaffer system) and by
Evans in his catalogues of the American Hesperiidae.
THE COMSTOCK-NEEDHAM SYSTEM
The Comstock-Needham system (Fig. 4) is followed by most
present-day American writers, although there are modifications
of it utilized by one or another. This scheme is based on the
venation of all insects, not just Lepidoptera, and thus has more
universal application than other systems. The most anterior
forewing vein is denoted the subcosta (Sc), the true costal vein
being lost in at least the butterflies, the next five are branches
44
LEE MILLER
/. Res. Lepid.
Fig. 4. — Venation and extradiscal cells of a hypothetical butterfly according
to the Comstock-Needham system. The symbols are explained in the text.
8(2):37~48, 1969(1970)
VEINS AND CELLS
45
of the radius (Ri, Rg, R3, R4 and Rg, from the anterior to the
posterior one), the next three are branches of the medius (from
the anterior, Mi, M2 and M3), the following two are branches of
the cubitus (from the anterior, Cui and Cug , or according to
some authors, Cui and Cui , respectively), and those veins
arising posteriad of the discal cell are the anal veins (A): the
one present in all butterflies is known as 2A (it also may be
denoted lA, depending on whether the fold posteriad of the
last cubital vein is considered the remnant of lA or of Cu2, and
this depending on the interpretation of the cubital veins). The
anal veins are also known as vannal veins, in which case they
are abbreviated IV, 2V, etc. The spur vein anteriad of the hind-
wing main veins (the precostal vein of other systems) is the
humeral vein (h), the anterior main vein may be considered as
the subcostal vein and the first radial branch (Sc+Ri), the
second main vein is the radial sector (the fusion of all of the
radials except Ri and abbreviated as Rs, not Rg as stated by some
authors), then come the three branches of the medius (from the
anterior to the posterior, Mi, M2 and M3), and the last two veins
arising from the cell are branches of the cubitus (again Cui or
Cui the anterior one and Cu2 or Cui the posterior one). The
anal (or vannal) veins arise posteriad of the cell and are de-
noted as lA (IV), usually absent in butterflies, 2A (2V) and
3A (3V) from the discal cell to the inner margin. The stalk
veins delimiting the discal cells of both wings are anteriorly the
radius (R) and posteriorly the cubitus (Cu). Many authors
still refer to the crossveins at the end of the discal cells as the
upper (udc), middle (mdc) and lower (Idc) discocellulars, but
the system is explicit in that these crossveins are named for the
veins they connect, so that the crossvein between Mi and M2 is
denoted mi-ms. Note that the initials are in lower case in this
instance.
There are at least two methods of designating the extra-discal
cells in the current literature: the discal cell (D) is the same in
both. Both Klots and the Ehrlich in their books on North Ameri-
can butterflies name these cells for the veins forming their
anterior boundaries (Fig. 4), so that the cell bounded by veins
Ml and M2 is cell Mi. Other authors use a system of naming
these spaces similar to that proposed by Rothschild and Jordan
(Fig. 5), using the names of both boundary veins to designate
a cell; thus, the cell between veins Mi and M2 is space
M1-M2 (note that in this instance the symbols are capitalized
to avoid confusion with the terminology for crossveins).
46
LEE MILLER
/. Res. Lepid.
Fig. 5. — -Wings of a hypothetical butterfly showing the proposed uniform
system of nomenclature for veins and extradiscal cells. The symbols are
explained in the text.
8(2):37-48, 1969(1970)
VEINS AND CELLS
47
SPECIAL STRUCTURES
In the discussion of the various systems I have mentioned the
small spur vein at the anterior basal part of the hindwing, the
precostal vein (pc, pcv) of older systems or the humeral vein
(h) of the Comstock-Needham system. Zeuner (1943, Ann. Mag.
Nat. Hist., 11/10: 290) considered that this vestigial vein repre-
sented either the costa (C) or the first branch of the primitive
subcosta (Sci), but other authors have not been certain or have
considered that this vein is unrelated to the main veins and
arose de novo in Lepidoptera.
In a few groups of butterflies, for example, the Brassolinae the
proximal part of the anterior main vein of the hindwing is
divided into two members (Fig. 5). There is no provision in any
of the systems to name these two veins, except the Comstock-
Needham system where the anterior member is Sc and the
posterior Ri. The more or less triangular cell formed by these
two veins and the anterior boundary of the discal cell is called
the precostal, prediscoidal or simply the basal cell in most
systems, but may be designated as cell Sc-Ri in that persuasion
of the Comstock-Needham system advocating the naming of the
cells for the veins bounding them.
DISCUSSION
The Herrich-Schaffer, "Indian” and Rothschild-Jordan systems
simply are not applicable in view of modern evidence as to the
identity of veins. Since these systems were based on Lepidoptera
only, they are not applicable for other insect groups. The syn-
thetic “English” system is not only inapplicable to other groups
of insects but also is not completely reliable for Lepidoptera.
This system was devised primarily for butterflies and is singularly
fitted only for them, but the scheme may fail when applied to
some moth groups that have more or fewer veins. This system is
in wide use in the moths, largely because it was employed by
pioneer Heterocera workers such as Meyrick and Hampson, but
at least the latter author had problems in applying the system
uniformly throughout his work. A system, then, to be most
valuable must offer the opportunity to draw homologies be-
tween the venation patterns of diffuse groups.
The remaining system, the Comstock-Needham system, is the
only relatively natural one that can be used not only for Lepidop-
tera but also for other groups of insects, and, as such, is the
48
LEE MILLER
/. Res. Lepid.
most useful to entomologists. I will not address myself to the
problems of cubital and anal vein nomenclature; these are
matters for individual workers to decide ( which scheme is being
employed soon becomes evident from reading a paper, anyway).
The use of the Comstock-Needham system enables anyone,
lepidopterist or not, to know just what vein is being referred to
in a paper on Lepidoptera. Since lepidopterists are also entomol-
ogists, and since the Comstock-Needham system is the system
that is recognized by entemologists of all specialties, it would be
best if lepidopterists adopted that system used by the great
majority of other entomologists.
The nomenclature of the extradiscal cells is somewhat more
difficult. The scheme promoted by Klots and the Ehrlich of
naming these cells for the veins anteriad of them is in direct
opposition to the “English” system in which the cells are named
for the veins posteriad, and a person familiar with one system
will almost invariably misinterpret the other. A non-lepidopterist
will incorrectly interpret such notations half the time, if indeed,
he can decipher the numerical system at all. By contrast, naming
the cells for the veins bounding them removes any confusion as
to just what cell is under discussion. This rationale is not new,
having originated with Rothschild and Jordan before the turn
of the century, but this idea has been more or less ignored
recently. However, because of its absolute clarity it seems the
best solution to the problem of accurately denoting cells.
An example of this preferred system for naming veins and cells
is given in Fig. 5.
ACKNOWLEDGMENTS
I would like to thank my wife, Jacqueline, A. C. Allyn and
Dr. E. D. Cashatt for reading this manuscript and making certain
suggestions on it, many of which have been incorporated into
the final draft.
Journal of Research on the Lepidoptera
8(2):49=50, 1969(1970)
1160 W. Orange Grove Ave.p Arcadia, California, U.S.A. 91006
© Copyright 1969
NOTES ON
LARVA AND HABITAT OF
CALLOPHRYS FOTIS BAYENSIS
(LYCAENIDAE)
RICHARD M. RROWN
1385 Palm Ave., Martinez, Calif. 94553
Guppy (1959) states that Callophrys fotis mossii (H. Edw. )
has been feeding long enough on its host to evolve a ‘'compli-
cated system of protective coloration/’ This is also true with
Callophrys fotis hayensis (Brown). The eggs hatch in three to
five days (in captivity) after being laid on the under side of
leaves on the flower stock of Sedum spathulifolium (Hooker).
The larva are green when they hatch; they remain this color,
if they continue to feed on green leaves, but if they feed on
older red leaves the caterpillars are red or pink. In the last
instars, if they feed on the flowers which are yellow, the cater-
pillars then turn yellow. The pupa is brown with dark specks
and a light pubescent covering.
The young and middle instars feed by boring into the thick
succulent leaves and by eating the insides; many times the
only thing one sees is a pile of wet frass.
In captivity the larvae which feed entirely on flowers are
healthier and mature faster than larvae which feed entirely on
leaves.
With the staggering rate at which natural areas are being
destroyed, it is good to have pictures published of this en-
dangered habitat. A fairly thorough description of the San
Bruno Mountains, San Mateo Co., is given by McClintock, and
Knight (1968), “A Flora of the San Bruno Mountains, San Mateo
County, California.” The habitat is an area without trees and
49
50
RICHARD M. BROWN
with low growing vegetation, the tallest being approximately
three feet. This area gets no protection from wind and fog
from the Pacific Ocean.
Fig. 1. — Western Side of the San Bruno Mountains, San Mateo County.
Fig. 2. — North facing slope of canyon below radio towers.
Fig. 3. — Sedum spathulifolium (Hooker), small, low growing, flowers ap-
proximately six inches tall.
Journal of Research on the Lepidoptera
8(2):51-52, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
A POSSIBLE NEW HYBRID COPPER
CHARLES R. CROWE
5027 N.E. 23rd Am.
Portland, Oregon
Fig. 1.
Dorsal (upper) and ventral (lower) views of male Lycaena rubidus (left),
editha ( right ) , and the hybrid form rubidus X editha ( center ) . Photography
by Don Fames, Portland State College.
51
52
CROWE
An interesting curiosity appeared this season near Burns,
Oregon that should be of interest to students of Lycaena. It is
and apparent hybrid Lycaena ruhidus X editha. To my knowl-
edge this hybrid has apparently been unreported in nature to
date. It is to be retained in the collection of the author.
The speciment was taken in company of normal forms of both
L. ruhidus and editha at Devine Canyon, 6 VIII 65, Highway
395, twelve air miles NNE of Burns, Harney Counyt, Oregon at
4,800 feet. The canyon is a primarily pine- juniper area that is
surrounded by sagebrush, and associated with a wet meadow
along Theimmer Creek that is lined with willow, birch, and
aspen. From this locality are also known to occur L. heteronea,
cuprus, and helloides.
As can be seen in the photographs, the main distinctions of
the hybrid are based on four points; the intermediate nature
of the dorsal ground color, the ventral HW pattern, the outline
of the FW, and invasion on the DHW anal margin of spots
typical of the editha pattern. Comparison specimens of L. editha
and ruhidus have been pictured also.
Journal of Research on the Lepidoptera
8(2):53-54, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
CONTROLLED ENVIRONMENT EXPERIMENTS
WITH PRECIS OCTAVIA CRAM.
(NYMPHALIDAE)
L. McLEOD, B.Sc., F.R.E.S.
25 Sleford Close, Balsham, Cambridgeshire, England
Continued from:
Journal of Research on the Lepidoptera,
volume 7(1) :18.
COLOR PLATE
Larvae and pupa
(adults to come in future issue)
53
54
McLEOD
Precis octavia sesamus
‘"plain” larva, 24° C
“striped” larva, 24° C
plain” larva, 30 °C
gold pupa
Journal of Research on the Lepidoptera
8(2):55-64, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
ON THE GUNDER COLLECTION
OF ARGYNNIDS
( Lepidoptera : Nymphalidae )
L. P. GREY
Rt. 1, Box 216, Lincoln Maine, U.S.A. 04457
In contemporary butterfly literature, F. Martin Brown’s
numerous papers on earlier collectors and collections have been
especially valuable. It is well that students should be reminded
of the importance of historical background and especially good
that they should learn of the extent to which nomenclature is
based on the art of “second guessing”. Indeed, it may be
uncomfortably close to the truth to say that the majority of taxa
proposed for butterflies prior to the twentieth century now rest
and must be allowed to rest on the deductions of specialists
concerning what might be termed accidents of history. The
identification of syntypical specimens and the selection from
them of suitable lectotypes has been a major preoccupation
of revisionary authors, with no end yet in sight. And what
fascinating snarls have been revealed, when digging to bedrock
for “origins”! Occasionally, even the apocryphal rumors and
gargantuan tales of the earlier giants have to be given some
weight when tracing material, as witness the stories of Herman
Strecker’s high silk hat.
It becomes painfully clear how large is the role of historical
happenstance in shaping nomenclature when it is recalled in
terms of concrete examples. A classic one of course is the
handling of the W. H. Edwards collection. How often students
find themselves wishing that this material could have been
preserved exactly as Edwards arranged and labeled it when
it was in his hands, at Coalburg!
Which reminds me that in a small way I was involved in an
55
56
L. P. GREY
/. Res. Lepid.
analogous history. Therefore I believe it is a duty to recount
what I can remember of the handling of the J. D. Gunder series
of argynnid butterflies after they came to The American Museum
of Natural History in New York.
A number of things were evident at that time, merely from
Gunder’s personal arrangement of his specimens. These were
details which became obscured or lost when the series were
shuffled from their original ordering and incorporated into the
Museum drawers. I recall in particular several oddities of
interest to Speyeria students. For one thing, the taxonomic
status of Argynnis pfoiitsi Gunder (1933, p. 171) appeared to
me then in a light which no future reviser ever could be ex-
pected to apprehend, as I shall explain. Also, a few questions
were raised which to this day remain unclarified.
As a visiting guest I had no part in policy making, but as
a bystander I was impressed by the solicitude of Michener,
Klots and dos Passos as they discussed how best to conserve
the values and potential in the Gunder material. It was an
amusing bylight, too, I thought, that Lutz, who at that time
was chairman of the Department, seemed to be indiflerent to
the whole affair. His passion was for experimentation, probing
the physiological and other biological attributes of insects; one
might say he was very modern in his contempt for the mere
“collecting” of dead butterflies.
The burden of guiding decisions thus fell mainly on dos Passos
and Klots. It must not be assumed that they were insensitive
to the value of the Gunder Collection purely as an historical
monument. It is doubtful if any of our students who are under
fifty or sixty years of age can really sympathize with their
dilemma. One would have to be able to evoke the historical
“then” and one would have to have lived through the period
to know and understand just how incredibly far the study of
Nearctic butterflies has progressed since that relatively short
time ago.
In retrospect, this seems to have been a turning point.
Butterfly classification had evolved mainly from odds and ends,
and even singletons, acquired at random as chance had afforded,
usually bearing such edifying labels as, e.g., “Oregon Territory”.
The people then recognized as specialists and “best authorities”
were laboring under a handicap beginning to be felt but im-
possible to overcome. Geographically representative series
8(2h55-64, 1969(1970) GUNDER ARGYNNIDS
57
simply did not exist. There was no possibility of examining
region-to-region intergradings and discontinuities, much less to
examine them for sympatrisms or to spin theories of their
correlations with late Pleistocene refuging and ecogeographic
factors.
Gunder may not have been the first to realize this need, but
it can be emphasized that he was certainly the leader at the
time, in this field of attempting large-scale geographic cover-
age. It was his vision and industry which Klots and dos Passes
determined to carry forward and amplify. Nobody should fault
them for scattering these particular bones of history. They
broke up Gunder’s arrangements, true, but only to lay the
foundations of one of the great study collections of North
American butterflies. I think they did the right thing; I lived
in the era, too, and can remember how imperative our needs
were, for better coverage, for continental surveys.
Reasonable care was exercised to keep the material in order.
Every specimen was ticketed to identify its derivation from
the Gunder Collection. Specialists were consulted before the
plaques were opened and their contents dispersed. The fact
of the specimens being in the the book-type Riker Mount cases,
on cotton, was of course one of the major factors prompting the
decision to rework Gunder’s material. The papered excess
was spread for later incorporation. The type specimens of taxa
authored by Gunder were taken into the Museum’s type col-
lection which is maintained separately and given special care.
This left the plaques, which Gunder considered to be his col-
lection proper.
Dr. dos Passes invited me to help him pin and reclassify the
“Argynnis\ for three reasons: (1) We were then planning a
jointly authored revision of the Nearctic species of these
butterflies. (2) And prerequisite to this we had to rearrange
and make usable the then-chaotic Museum collection, incor-
porating with it the extensive Gunder series. (3) Also, it seemed
desirable that we should share responsibility of preserving
whatever taxonomic or other data or deductions might appear
from the original plaque arrangements and sortings. As I
recall, we spent something like ten full working days merely
to shuffle to a “species-by-States” arrangement, before any
“study” could be possible. Incidentally, a recent (1969) check
indicates that the geographic order has been maintained despite
considerable additions. Students who go here and are given
58
L. P. GREY
J. Res. Lepid.
instant access to whatever may be available of particular series
from particular areas should realize the debt they owe to people
like Gunder, Klots, dos Passes, and to the present Curator, Dr.
Rindge, who keeps the series in scrupulous order and has added
largely from his own field collecting.
Even so, and with all the work which has been done to ease
the labors of researchers, I had a unique and never-again oppor-
tunity to see things which are now beyond recall. I saw pre-
cisely how Jeane Gunder interpreted taxa and categories, and
I think it is long past time that somebody should speak up and
defend his abilities. He seems to be remembered principally
as a trifler with “aberrations”, an arch-splitter. Few students
seem to have any idea of his true dimensions as a pioneer. I
noted instance after instance wherein he had lumped or juxta-
posed taxa then rated as separate entities. The sheer size of
his accumulations witnessed more eloquently than he, himself,
ever managed to explain, of the importance he attached to geo-
graphical variation and of his concern to extend coverage to
include generous population samples from as many localities as
possible. Eastern lepidopterists, seeing this collection for the
first time, were introduced to butterfly study in a new dimen-
sion; it was quite a jolt to some of them who- had dismissed
Gunder as a wild amateur.
I can testify for the argynnids that Gunder s arrangements
bespoke not only his appreciation of the basic needs for ex-
tensive comparisons, but also a great deal of research in the
literature and the study of preserved type series. His taxon
usages in the main were up to present standards but naturally
some of his ideas of “species” now seem outmoded. There is no
need to eulogize him unduly; he made his share of blunders,
and misdeterminations, and, as amply proclaimed by his critics,
he wasted a disproportionate amount of energy in futile at-
tempts to give nomenclatorial status to aberrations and minor
color forms.
On balance, however, he surely deserves more credit and ap-
preciation than seems to have been accorded him as one of
our leading authors. The labels he put under his collection
series I would say revealed a better grasp of identities and
entities than can be claimed for any argynnid student prior to
his time. They resulted, I am sure, from painstaking study
combined with a really formidable taxonomic intuition. As
for his blunders, one suspects that future workers will find
8(2):55~64, 1969(1970) GUNDER ARGYNNIDS
59
that “me and thee” also have sinned: it is impossible to work
through any large collection without coming across the occa-
sional lapses from virtue such as happen to us all. It is hardly
fair to charge the man with errors which were, so to speak,
inherent and embalmed in the listings and concepts of his day;
leaving these aside only the few mistakes detailed in following
paragraphs were noted, to which will be added my personal
appraisal of their historical origins.
Gunder has been charged with one major taxon-error, namely,
his misapprehension of Argynnis platina Skinner (1897, p. 154).
I was in a position to understand how this error arose, since
I had visited the Academy of Sciences and had studied Skinner’s
Utah material, shortly after Gunder had been there for the same
purpose. Thus, I am safe in presuming that Gunder saw* exactly
what I did, in the way of Skinner-labeled material. It thus
seems evident that he merely accepted, on Skinner’s authority,
that the variation range in Skinner’s ‘^platina” included forms
which we now relegate to another species. It may as well be
admitted that Skinner’s legacy is a confused one; he apparently
was unable to separate his own ^‘platina’ from his own utah-
ensis (1919, p. 216). I recognized that his series were badly
mixed and had the good fortune to be able to check my con-
cepts with Nabokov; the latter had been collecting in Utah,
had a good eye for species discrimination, and had been looking
into these questions through spot-locality comparisons of
sympatrisms, extent of local variation, etc. We agreed that
Skinner never did learn to separate the Utah argynnids.
But Gunder tripped over Skinner’s mistakes, with the result
that he took away the impression of '^platina' as applying to
‘"utahensis” . Then, in a very interesting display of taxonomic
virtuosity, Gunder thereafter consistently applied '‘platina' in
the erroneous way he had apprehended. Thus it came about
that Gunder ’s plaque of "platina' was filled with Idaho greenish-
disk egleis (Behr) (“1863”: 1862, p. 174) of the sort which dos
Bassos and I later dubbed "linda". Knowing this much of the
story it is clear that Gunder would assume one of the major
elements in Utah argynnid variation to be nameless. Hence,
his description of "pfoutsf, justifiable by all that he had been
able to learn of types and of natural populations.
In this instance one sees again the prime importance of back-
ground data when assessing nomenclature. Granting the above
bylights on "pfoutsf an adjudication of its status follows inevi-
60
L. P. GREY
/. Res. Lepid.
tably: it drops to synonymy, naturally, but it should be of in-
terest also to know that it does not represent mere ignorance
or a propensity to split, on Gunder’s part, but is rather a wholly
excusable mistake with a logical historical cause.
Another incongruity in the Gunder series, one I have kept in
mind over the years, was the occurrence in Nevada-labeled
material (Clark and Lincoln Counties, leg. Eugene Schiffel) of
specimens obviously representing subspecies of atlantis (Ed-
wards) (“1862”: 1863, p. 54) and of hydaspe (Boisduval) (1869,
p. 60). These were of facies suggesting derivation from Mon-
tana or perhaps British Columbia. The geographic association
seemed rather weird, even then, at a time when very little was
known of distribution. Even today it might be risky to aver
precisely what does or does not occur around the Spring Moun-
tains area in the way of Speijerui. However, from everything
presently witnessed and conceived, this bears the earmarks of
some preparator’s mistakes. I mention it to ease the minds of
investigators who may run across these specimens in the Mu-
seum. It seems best agreed that whoever will accept these
records as authentic should bear the burden of proof. Quite
likely they resulted from some scrambling of envelope data
but at any event this probable boo-boo involves merely the
geographical labeling, and not taxon confusion. But another
incongruity I took note of seems to involve a little of both, who
knows?
The plaque of Argynnis whitehousei Gunder (1932, p. 279)
consisted of 3 males and 5 females identifiable as an aphrodite
(Fabricius) (1187, p. 62). But in the same plaque were 8 males
of an egleis subspecies (my identification). These latter were
in a facies which would have been tolerable if they had been
labeled as from “Utah”, instead of as from “British Columbia”.
They were doubly suspect to me also since I did not know then
and still do not know of any authentic British Columbia records
of egleis, this being a species which seems to taper off to rarity
in northern Montana.
I offer no guaranteed solution to this strange action of
Gunder’s; probably it is best to treat it as an unresolved mystery,
which, in any event, is the present state of knowledge re
northernmost distribution of egleis. Still, it is tempting to
express my suspicions, since they might provide another lead
in case that Canadian students should fail to find egleis after
due search in the indicated region: I can vouch for the fact
8(2):55-64, 1969(1970)
GUNDER ARGYNNIDS
61
that Tom Spalding supplied Gunder with some material; I
learned this from my correspondence with both of them. From
the appearance of these specimens in question I have reason
to guess that they might have derived from the Provo region
of Utah, which Gunder’s involvement with Spalding would
rationalize. But as it stands they are purportedly from Jaffray,
B. C., August 1-5, 1929, leg. Whitehouse. It is very definite,
then, that Gunder was guilty here of one of his rare lapses, mak-
ing that most embarrassing of all taxonomic mistakes, namely,
confusing things distinct in nature. To top it off, I fear he had
another visit from the scramble-gremlin which misplaces geo-
graphical labels on spreading boards. At least, the question
must be answered: What actually does occur in the vicinity of
Jaffray, in the way of an egleis subspecies?
In summary, then, many values were lost when Gunder’s
"‘Argynnw* were removed from their plaques. Today undoubt-
edly we would photograph them before tampering. The fact
remains, however, that these specimens, vastly enriched by later
additions, have served the true purpose intended by Gunder
and still remain fully accessible to interested students in the
precise but expanded concept and vision of Gunder, which was
to build toward a total view of North American butterfly
speciation and subspeciation. Unfortunately, the thing which
was lost in the process was an intangible vignette of Gunder
himself, as reflected by his handwork.
At this late day, the only amend possible is to aflBrm for what
my personal opinion may be worth that Gunder had rare
natural talent as a taxonomist despite popular impressions to
the contrary. I had the privilege of seeing for myself that his
competence in sorting argynnids was quite amazing; very few
students even today can approach his abilities in this depart-
ment. And those who can remember what it was like, back in
that quite recent and yet curiously remote era, to confront
Western Speyeria en masse — we, at least, know very well the
debt we owe to Gunder.
Merely from his sortings, innumerable instances could be
cited of his acuity. Referring back to the blunder in the lohite-
housei plaque for example, one still could note how unerringly
he had fingered out the aphrodite variation in the remainder of
his British Columbia material, even from localities where
aphrodite runs excruciatingly parallel to other species. In this
and in many other instances of an analogous nature, his accurate
62
L. P. GREY
/. Res. Lepid.
separations of parallel sympatres have stood unmodified over
the years in the face of inspection by students with far larger
data than ever were available to Gunder. He stood unmatched
among his contemporaries; he was a far better argynnid taxono-
mist than McDunnough, for example, as can be seen from the
historical record: McDunnough blundered seriously with some
of the Western Canada parallels, even to the extent of assem-
bling a mixture of entities in type series! My personal debt to
Gunder is no small one. Dr. dos Passes and I became heirs to
all of his extensive preliminary work with argynnids, and, as
it has turned out, could have found no better source of properly
sorted and correctly identified material.
Additionally, students should bear in mind that Gunder ’s
approach to difficult genera was altogether modern although
practically new and unheard of at the time. He first assembled
huge and geographically representative material, which he
attempted to sort out to “species”, with a shrewd eye to sym-
patrisms and to variation as correlated with geographic barriers
and opportunities for dispersals. In the case of the genus
Euphydryas he went even further, to synthesize all of these
facts with the evidences of genitalic structures. Indeed, his
1929 revision of the latter genus remains to the present day
one of the landmark papers which have shaped our modern
classification and concepts of butterfly species.
Given more time, it is altogether probable that Gunder would
have revised ‘‘Argynnis” along the identical lines followed by
dos Pass os and myself. It was clear that he was quite far along
in the data-gathering stage, and that he would have made short
work of the niney to a hundred and twenty-five or so “local
species” then cluttering our lists and manuals. It cannot be
repeated often enough that this man was not a splitter. He was
a synthesizer, born before his time. We had no difficulty in
following his ideas as expressed by his collection arrangements,
and found relatively few puzzles and contradictions other than
those described herein which seem mostly due to scrambled
data. So, for argynnids, what with Gunder’s published descrip-
tions and the careful preservation of his specimens at the
American Museum, there is little for future historians to stumble
over, it would appear, if they will steer away from the super-
ficial and altogether false presentation of Gunder as a playboy
amateur.
Among other misfortunes which dogged Gunder, there remains
8(2):55^64, 1969(1970) GUNDER ARGYNNIDS
63
a major canard which seems to pass unanswered. Speculations
continue to circulate that he did not do the work on Euphydryas^
that it may have been the product of a hired collaborator. Be-
fore the obscurity of years closes over this latter revision it is a
matter of urgency that any of the older generation having
recollections or letters bearing on this subject should publish
them. This is clearly an instance wherein '"trivialities” might
prove to have major historical importance. One fact seems
assured: None of the original dissections or drawings were in-
cluded in the material purchased by the Museum.
I know of no helpful data which might apply to this riddle.
I exchanged relatively few letters with Gunder and in them
there were no mentions of genitalic studies and only a few
references to Western Euphydryas problems. Therefore, my
personal curiosity, and I am sure the curiosity of other students,
remains unsatisfied. We have a natural desire to know some-
thing of Gunder’s methodology, of the material he assembled
and his understanding of it. We have a duty to future research-
ers who will be equally curious. But as it stands, our estimation
of the 1929 Euphydryas revision as a brilliant achievement seems
best enforced by the fact that nobody seems able, even with
vastly expanded material and knowledge, to come up with a
better synthesis.
Are we never to learn more of the background of this mys-
terious feat? Perhaps, then, I should make bold to offer a
comment which may have some incidental bearing on it. I
know nothing about Euphydryas, but I did see how splendidly
Gunder was brushing through the utter confusion which then
prevailed in argynnids. From that experience I know that
Gunder had an innate gift, a brilliance denied or only grudgingly
recognized by his critics. Thus, I would be willing to defend the
idea that Jeane Gunder needed no hired talent to supplement
his own genius. Whoever can tell us more should do so.
REFERENCES
BEHR, HANS HERMAN, “1863” (1862). On Californian Argynnids.
Proc. Calif. Acad. Nat. Sci., 2: 172-177.
BOISDUVAL, JEAN BAPTISTE ALPHONSE DECHAUFFOUR DE, 1869.
Lepidopteres de la Californie. Ann. Soc. Ent. Belgique, 12: 1-28, 37-94.
EDWARDS, WILLIAM HENRY “1862” (1863). Descriptions of certain
species of diurnal lepidoptera found within the limits of the United
States and British America. Proc. Acad. Nat. Sci. Philadelphia, 14:
54-58.
64
L. P. GREY
/. Res, Lepid.
FABRICIUS, JOHANN CHRISTIAN 1787. Mantissa insectorum, sistens
eorum species nuper detectas adiectis characteribus genericis, differ-
entiis specificis, emendationibus, observationibus. Copenhagen, Christ.
Gotti. Proft. 2: 1-382.
GUNDER, JEANE DANIEL 1929. The genus Euphydryas Scud, of Boreal
America ( Lepidoptera Nyinphalidae). Pan. Pac. Ent. 6: 1-8, 10 pis,
3 maps, 1 table.
1932. New Rhopalocera (Lepidoptera). Canadian Ent., 64: 276-
284.
1933. Additional New Rhopalocera (Lepidoptera). Can. Ent.
65: 171-173.
SKINNER, HENRY 1897. Notes on Rhopalocera, with descriptions of new
species and varieties. Canadian Ent., 29: 154-156.
1919. A new species of Argynnis from Utah. (Lepid., Rhop. ).
Ent. News, 30: 216.
Journal of Research on the Lepidoptera
8(2):65-=68, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
ESTIMATION OF NATURAL MUTATION RATES
FOR ALBINISM IN TWO SPECIES OF THE
SATYRID GENUS CERCYONIS
THOMAS C. EMMEL
Department of Zoology, University of Florida, Gainesville 32601
In the past decade of field work and laboatory research with
the woodnymphs of the Nearctic genus Cercyonis (Satyridae),
I have had the opportunity to collect data on the frequency of
various mutant genes within the four species: C. pegala, oetus,
sthenele, and meadi (See Emmel, 1969, for taxonomic summary).
Fig. 1-2. — ^Dorsal surfaces of albinic and normal male specimens of Cercy-
onis oetus from a population in the Reese River Valley, Lander County,
central Nevada.
Fig. 3-4. — Ventral surfaces of same specimens.
65
1
66
T. C. EMMEL
/. Res. Lepid.
Fig. 5-6. — Dorsal and ventral surfaces of partially albinic female specimen
of Cercyonis pegala from Boardman, Morrow County, Oregon.
8(2):65~68, 1969(1970)
MUTATION RATES
67
An additional point is that spontaneous mutant specimens are
probably eliminated from the populations largely by mate se-
lection. Brown females undoubtedly refuse strange-looking
white males (e.g., see Sheppard, 1961) and brown males most
likely fail to approach white females as being of the "wrong”
species.
I. Cercyonis oetm
On July 12, 1969, a totally albinic male C. oetm was taken by
John F. Emmel in a population of brown specimens located four
miles northeast of the Reese River on Highway 2, 5700 feet ele-
vation, west-southwest of Austin, Lander County, Nevada. The
dorsal and ventral surfaces of this specimen are compared with
the normal male phenotype of this population (itself being
extraordinary; see Emmel and Emmel 1970) in Figures 1-4.
The only significant departure from complete lack of pigment is
in the forewing ocelli, which are light brown instead of the
usual black. The specimen is in essentially freshly emerged
condition.
II. Cercyonis pegah
A partially albinic female individual of Cercyonis pegala was
collected on the west side of the town of Boardman, 200 feet
elevation, in Morrow County, Oregon, on July 11, 1964, by
Edwin M. Perkins and Stephen F. Perkins. In this specimen,
the albinic portions are mainly restricted to the outer half of
each wing (but both surfaces).
DISCUSSION
One can calculate an approximate rate of spontaneous mu-
tation for the expression of albinism by dividing the number
of known mutant indivduals by the total number of individuals
observed. I have personally examined or seen in the field more
than 12,000 individuals of C. oetm and more than 6,000 indi-
viduals of C. pegala, at a conservative estimate. With respect
to albinism, the two specimens reported here are the only
mutants I have seen. A number of other lepidopterists with
many years in the field confirm these observations, adding still
more to the base number observed for each species.
Thus we can estimate the probable maximum natural muta-
tion rate for albinism in the two species, within an order of
magnitude, as:
Cercyonis oetm lO-® (.00001)
Cercyonis pegala 10® (.00001)
These figures, of 10-® per gene per generation, are in the same
68
T. C. EMMEL
J. Res. Lepid.
order as those known for Drosophila and man (Dobzhansky,
1951, p. 59) and for the domesticated silkworm, Bombyx mori
(Tazima, 1964, p. 179-180).
Only two mutations involving albinism have come to my at-
tention, and the purpose of this note is to provide an estimation
for the spontaneous rate of mutation for this character in two
species of these satyrids.
LITERATURE CITED
DOBZHANSKY, THEODOSIUS, (1951). Genetics and the Origin of Spe-
cies. Columbia University Press, New York. 364 pp.
EMMEL, THOMAS C., (1969). Taxonomy, distribution and biology of the
genus Cercyonh (Satyridae). I. Characteristics of the genus. Journ.
Lepid. Soc., 23:165-175.
EMMEL, THOMAS C., and JOHN F. EMMEL, (1970). An extraordinary
new subspecies of Cercyonis oetus ( Lepidoptera, Satyridae) from cen-
tral Nevada. Pan-Pacific Entomologist, in press.
SHEPPARD, P. M., (1961). Some contributions to population genetics
resulting from the study of the Lepidoptera. Advances in Genetics,
10:165-216.
TAZIMA, YATARO, (1964). The Genetics of the Silkworm, Academic
Press, London. 253 pp.
BOOKS;
NOTICES
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastimil Choc. Paul Hainlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophieal
Library, N.Y.
WANTED:
Brephidium exilis, B. fea, B. isophthalma. Life material and speeimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, Galifornia 91006,
IN PREPARATION:
BUTTERFLIES OF NORTH AMERIGA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systematics, biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
Manuscripts for immediate publication in this JOURNAL. With color
lAay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3x5 card in the form desired and they will be printed in the
next following is.siie of the JOURNAL.
THE JOUI^NJAL ©F RESEARCH
ONI THE LEFIJD0FTERA
Volume 8 Number 2 June, 1969
IN THIS ISSUE
Nomenclature of Wing Veins and Cells
Lee D. Miller
37
Larva and Habitat of CaUophrys fotis bayensis
R. M. Brown
49
A possible new hybrid copper
Charles R. Crowe
51
Controlled environment experiments with
Precis octavia C. (color plate) L. McLeod
53
On the Gunder collection
L. P. Grey
55
Estimation of natural mutation rates for albinism
in two species of the Satyrid genus Cercyonis
T. C. Emmel 65
THE JOURNAL OF RESEARCH
©NJ THE LERIJDORTERA
published by
The Lepidnptera Research Foundation, Inc.
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Journal of Research on the Lepidoptera
8(3):69-90, 1969(1970)
1160 W. Orange Grove Ave.^ Arcadia, California, U.S.A. 91006
© Copyright 1969
DEVELOPMENT OF THE MARKINGS
ON THE PUPAL WING OF
PIERIS RAPAE (PIERIDAE)
JOHN M. KOLYER
55 Chimney Ridge Drive, Convent, New Jersey, U.S.A.
INTRODUCTION
The development of the black markings of Pieris hrassicae
(L.) was studied by Onslow (1916), who reported that the
pupal wing became black all over when soaked in tyrosine solu-»
tion while selective darkening of the markings occurred in
tyrosinase solution. (The action of an oxidase, eg. tyrosinase,
on a colorless chromogen, e.g. tyrosine, is known to give melanin
pigments.) The conclusion was that the chromogen first was
deposited in the areas destined to become black, then oxidase
was supplied by the hemolymph, and finally atmospheric oxygen
reached the surface of the wing and caused darkening where
chromogen was concentrated. This explanation was modeled
after that of Gortner (1911a) for development of the color
pattern on the elytra of the Colorado potato beetle.
In contrast to Onslow’s result, Braun (1939) found that
'artificial pigmentation”, i.e. selective darkening of the markings,
occurred when pupal wings of Papilio ajax (L. ) and Ephestia
kuhniella (Zeller) were soaked in tyrosine solution. Braun’s
explanation was that the darkened scales were less chitinized
due to slower development and so were able to absorb the tyro-
sine solution. His conclusion was that oxidase is present in the
scales in general, but at the “certain time of development pig-
ment is present in the body” only the soft, less chitinized scales
are able to accept this “pigment” (chromogen).
The general problem of development of the wing pattern in
Lepidoptera has received study by Kuhn, Goldschmidt, Kohler,
and others, as reviewed in detail by Caspar! (1941). Brief
summaries are given by Wigglesworth (1965: 78) and Boden-
stein (1953).
69
70
JOHN M. KOLYER
/. Res. Lepid.
PLATE I
Fig. 1 (upper left) — Upper spot of wing from pupa ( 9 ) about 175
hours old, showing “relief” effect. The illuminating light beam was 10°
from the horizontal to give shadowing.
Fig. 2 (upper right) — Upper spot of wing from pupa ( 9 ) about 195
hours old ,witli markings in early stage of darkening. Lighting as in Fig. 1.
Fig. 3 (lower left) — Artificial pigmentation of wing from 190 hour old
pupa ( ^ ) by aqueous extract of hemolymph ( details in text ) .
Fig. 4 (lower right) — Artificial pigmentation of wing from 190 hour old
pupa ( ($ ) by undiluted hemolymph ( details in text ) .
8(3):69-90, 1969(1970)
WING MARKINGS
71
The object of the present work was to study pattern develop-
ment for Pieris rapae (L.) in order to contribute original obser-
vations as well as to evaluate the explanations of Onslow and
Braun.
EXPERIMENTAL
Source of pupae — Final instar larvae were supplied by the
U.S. Department of Agriculture (see Acknowledgement). These
had been reared on an artificial diet at 26 it 1°C and 45 it 5%
relative humidity under continuous cool white fluorescent light.
Development was completed on cabbage leaves from refrigerated
heads. Pupae were kept in a room at 70-80 °F and 28-35% rela-
tive humidity.
Dissection techniques. Early pupae, e.g. 90 hours old, pre-
sented more of a problem in removal of the forewing than did
pupae at later stages, after the wingcases had whitened at
about 135 hours. However, a successful procedure was to cut
off the head end of the pupa, just at the base of the wings, by
pushing downward with a razor blade. Then a small pair of
scissors was used to cut all around the wingcase. The wingcase
was placed on a table inside up, a small piece of blotting paper
was used to remove matter (including the hindwing) covering
the forewing, the nail of the left index finger was applied to
hold down the basal end of the integument, and the wing was
grasped at the base with pointed forceps and carefully peeled
off to be placed in water or aqueous solution.
As the time of eclosion drew near, it became possible to dis-
sect out the forewings merely by cutting the pupal case and
pulling the wing out by the base.
The wings shown in Plates I-III were allowed to dry in air,
and each was mounted on a microslide on a square of white
blotting paper under a cover glass, the latter being held in place
by a gummed label with appropriate hole. The photographs
were made through a lOOX microscope for Figures 1 and 2, a
lOOOX microscope for Figure 8, and a 16X microscope for the
other Figures.
Observations. — Most of the observations were made with a
stereo microscope at 16X. Illumination was a concentrated spot
of light from a microscope illuminator aimed down on the sub-
ject at a 45° angle. Features of the wing such as venation and
areas of translucent scales were seen most clearly against a
background of black felt, but judgment of degree of darkening
72
JOHN M. KOLYER
/. Res. Lepid.
PLATE II
Fig. 5 (upper left) — Wing from 160 hour old pupa blackened by ex-
posure to dopa solution followed by drying in air ( details in text ) .
Fig. 6 (upper right) — Artificial pigmentation of wing from 185 hour old
pupa ( 9 ) by dopa solution ( details in text ) .
Fig. 7 (lower left) — ■ Control wing (in water) for wing shown in Fig. 6.
Fig. 8 (lower right) — Dark scale from the spot of a wing from a 175
hour old pupa ( 9 ), artificially pigmented by dopa solution (details in
text). The scale was mounted in Permount (Fisher Scientific Co.) and
photographed by transmitted light through a microscope with lOX wide-field
ocular and lOOX achromatic objective (1.25 N.A., oil immersion).
8(3):69-90, 1969(1970)
WING MARKINGS
73
of scales was made against a white background. Close obser-
vation of the scales was made with a biological microscope at
lOOX or 430X with either reflected or transmitted light or at
lOOOX (oil immersion) with transmitted light.
Wings were soaked conveniently in solutions in uncovered
watchglasses at room temperature (70-80°F) for a few hours,
but longer times required closed containers to prevent evapor-
ation.
Solutions and reagents. —
Saturated tyrosine solution: Excess L-tyrosine (Matheson
Coleman and Bell) was shaken with deionized water. The con-
centration is reported to be 0.045% at 25°C (Anonymous, 1960).
Dopa solution (0.5%): In 20 ml deionized water was dissolved
0.10 grams of DL-3- ( 3, 4-dihydroxyphenyl ) alanine (practical
grade, Matheson Coleman and Bell). The solubility of the less-
soluble L-form is 0.5% (Anonymous, 1960). A 0.4% solution was
used by Gonnard and Svinareff (1951) as substrate for potato
tyrosinase.
Iodine reagent: According to the method of Campbell (1929),
a solution of 1.2 grams iodine and 1.6 grams potassium iodide in
1.5 ml water was added to 50 grams of 20% acetic acid to give a
clear, dark-red solution.
Tollens reagent: Small portions were prepared according to
Feigl (1954) and used immediately (the solution cannot be
stored as it decomposes and deposits explosive silver fulminate).
A convenient amount (about 0.6 ml) was given by adding 5
drops 10% sodium hydroxide to 5 drops 10% silver nitrate to give
a brown precipitate and then dissolving this by addition of 3
drops of a mixture of equal volumes cone, ammonium hydroxide
( 28-30% NHs ) and water.
Le Rosen formalin reagent: Since the reagent cannot be stored,
a small volume was prepared just before using by stirring 2
drops of 37% formaldehyde solution into 10 drops cone. (98%)
sulfuric acid in a watch glass. This is a variation (higher formalin
content) on the reagent according to Feigl (1954).
Misc. solutions: Concentrations are given in weight-%, e.g.
50% sulfuric acid was prepared by adding 50 grams acid to 50
grams water. The water used was always distilled and then
deionized. 4-Chlororesorcinol (Koppers Co.) was recrystallized
to give capillary melting point 108.5-110°C. The other organic
compounds were used as supplied by Matheson Coleman and
Bell or Eastman Organic Chemicals.
74
JOHN M. KOLYER
/. Res. Lepid.
11 12
PLATE III
Fig. 9 (upper left) — Wing from 175 hour old pupa ( 9 ) soaked in
oxygen-free dopa solution ( no pigmentation occurred ) and then allowed
to dry in air ( markings darkened; details in text ) .
Fig. 10 (upper right) — Wing from 189 hour old pupa ( $ ) tinted deep
pink by murexide fonnation ( details in text ) . The spot ( pale yellow ) is
faintly visible in the photograph.
Fig. 11 (lower left) — Wing dissected from pupa ( $ } with markings
just beginning to darken; shown after 18 hours in water-saturated air.
Fig. 12 (lower right) — Control wing (18 hours in water-saturated nitro-
gen) for the wing shown in Fig. 11.
8(3):69-90, 1969(1970)
WING MARKINGS
75
RESULTS AND DISCUSSION
1. Structural Changes During Development
Chronology of development. — The following notes on scale
development are preliminary observations based on a few dis-
sections. Times from pupation (final larval molt to give pupa)
are approximate and intended to be typical; some pupae devel-
oped more slowly.
27 hours: Careful dissection gave a tracheated wing with
frothy appearance at 430X. This seemed to consist of epithelium
with scales not yet grown. The wing epithelium of the freshly-
molted Pieris brassicae pupa is composed exclusively of stem
cells whieh later give the scale and socket arrangement (Lipp,
1957).
87 hours: Scales covered the wing. These were round, gener-
ally about 0.02 mm in diameter but some larger.
101 hours: Scales were generally round, approx. 0.04-0.09 mm
in diameter. Staining with 1% crystal violet (Colour Index No.
42555) in 95% ethanol or with a 1:1 mixture of saturated safranin
O (Colour Index No. 50240) in 95% ethanol with aniline water
(Shillaber, 1944) helped make the scales visible.
122 hours: Hairlike fringe scales on the margin were conspic-
uous against a black background. Scales on the wing were of
various shapes, some round, usually with a point, others some-
what elongated with three teeth. Length varied from about 0.04
to 0.14 mm. The impression was that some of the round scales
grow into the elongated form, length about 0.13-0.17 mm, that
is most common on the adult wing. Exposure to tyrosine or
dopa solutions darkened the scales and improved visibility at
100 or 430X.
135 hours: The wingcases became noticeably whitened to
the naked eye. Also noted was disappearance of the former
translucency of the pupa in the wing region when viewed from
the side against a source of light.
165 hours: The future markings (spots), though not at all
pigmented, were dimly visible through the pupal case.
175 hours: Female wings had a yellow appearance as viewed
through the pupal case.
195 hours: Darkening of the markings began. The most slow-
ly developing pupae reached this stage in 220 hours.
200 hours: Markings were completely darkened. The dark-
ening process required 4-5 hours at 80 °F. The wings shortly
later became hydrophobic whereas in earlier pupae they were
76
JOHN M. KOLYER
J. Res. Lepid.
easily wetted when dissected and placed in water.
220 hours: Eclosion.
Ages of pupae given below were adjusted in some cases in
order to indicate point of development in terms of the above
schedule.
Visible distinction of future-black scales. — The future-black
scales in the apical area and, particularly, in the spot(s) (one
on male, two on female wing) became noticeably different in
appearance (glossy by reflected light at a certain angle, trans-
lucent by transmitted light) at about 135 hours. A good de-
scription is “like spots of grease upon white paper” (Onslow,
1916). At 155 hours the scales all collapsed against the mem-
brane when the wing was dried in air, but at a later time, e.g.
175 hours, a “relief stage” (Braun, 1939) became obvious after
air-drying for only a few minutes. With side-lighting, the spot
scales appeared to have collapsed against the wing membrane
while the surrounding scales remained erected. An example is
shown in Plate I, Figure 1. This effect was no longer well-defined
on a pupal wing with markings just beginning to darken (Plate
I, Figure 2).
2. Artificial Pigmentation
Water and saline. — In no case among the many forewings
exposed to water did darkening occur in times up to 6 hours,
but at 37 hours (pupa 173 hours old) there was darkening
(brownish color) at the base and very slight darkening of the
future-black scales, the rest of the scales remaining the original
white. The other wing of the 173 hour old pupa was exposed
to 0.05 M sodium chloride for 37 hours with no darkening.
However, a wing from a 190 hour old pupa showed darkening at
the torn base in 0.05 M sodium chloride at 9 hours, and the spot
was very pale brown (wing itself very pale yellow-tan) at 48
hours. This NaCl concentration is in the general vicinity of the
chloride content of the pupal blood, e.g. 0.02 M in chloride for
the Pieris brassicae pupa ( Buck, 1953 ) . Onslow ( 1916 ) reported
slight darkening of markings on the pupal wing of P. brassicae
after 12 hours in “normal saline” (0.75% NaCl).
Hemolymph. — Hemolymph, as obtained in diluted form by
grinding pupae with chloroform water and filtering, was re-
ported by Onslow ( 1916 ) to give considerable darkening of the
markings of the pupal wing of P. brassicae in 12 hours.
In the present work, diluted hemolymph was prepared from
a 190 hour old pupa by grinding all but the forewings with 1.5
8(3);69-90, 1969(1970)
WING MARKINGS
77
ml deionized water and filtering to give a colorless, opalescent
liquid, in which one forewing was placed. The future-black
scales were very pale brown after 33 hours vs. no appreciable
darkening for the other (control) wing in deionized water. In
another experiment, the wings were removed from a 190 hour
old pupa, and the remainder of the pupa along with three pupae
with markings darkened was ground with 6 ml water (pH 5.6)
and filtered. Part of the filtrate (pH 6.6) was added to one
wing, and the remainder was adjusted to pH 8.0 with several
drops of 0.1% sodium carbonate solution and added to the other
wing. The same procedure then was repeated using chloroform-
saturated water. The result was that both water and chloroform
water extracts gave light-brown future-black scales visible against
the pale-tan future-white scales, but the water extract seemed to
give slightly more darkening (Plate I, Figure 3). Results at
pH 8.0 were not so good as at pH 6.6, especially for the chloro-
form water (negligible darkening of future-black scales). The
pH of P. rapae pupal blood has been reported as 5. 9-6.4 ( Buck,
1953).
A drop of clear, pale-green hemolymph was noted to exude
from the body of a 215 hour old pupa ( markings fully darkened )
from which the head end had been cut at the base of the
wings. This liquid was placed on one wing from a 190 hour
old pupa, and the other wing was placed in water as a control.
After 5 hours some darkening of the markings of the wing
with hemolymph was noted, and a small amount of water was
added to prevent desiccation. At 10 hours the markings were
well darkened in the hemolymph case (Plate I, Figure 4) vs.
no darkening of future-black scales for the control wing. The
contrast between markings and white scales was more pro-
nounced (white scales less darkened) for the wing shown in
Figure 4 than in artificial pigmentations with dopa solution.
Tyrosine. — Saturated tyrosine solution caused rapid blacken-
ing (in less than 30 minutes) at the edge of the torn base of
the wing, as did 0.5% dopa, presumably because of the tyrosinase-
containing hemolymph exposed in this area. For a 165 hour old
pupa the markings (spot and apex) darkened slowly; the scales
within the spot were pale gray after 6 hours. As a control, the
other forewing from the same pupa was soaked in deionized
water and showed no darkening after 6 hours.
Braun ( 1939 ) claimed that wings in tyrosine solution unfolded
(expanded), a phenomenon produced by '‘no other solution
tested”. In the present work there was much individual variation
in the extent of expansion, but all the aqueous chromogen solu-
78
JOHN M. KOLYER
/. Res. Lepid.
tions, 0.05 M NaCl, and deionized water itself gave this eflFect.
Using the distance from apex to outer angle (4-5 mm for un-
treated pupal wing, typically 14 mm in adult) as a measure of
expansion, the following values were noted for pairs of wings
from the same pupa: 11 mm for saturated tyrosine solution vs.
9 mm for water, 6.5 mm for 0.5% dopa solution vs. 7 mm for
water, and 10 mm for 0.1% sodium carbonate solution vs. 12.5
mm for water (the greatest expansion noted). Pupal age may
have a large influence on degree of expansion. Tyrosine solution
obviously is not unique in causing expansion, and possible c
marginal superiority over water or other aqueous solutions
would have to be demonstrated by a number of competitive
experiments.
Dopa. — The use of tyrosine solution soon was discontinued in
favor of 0.5% dopa, since the latter was found to be far superior
for artificial pigmentation. For example, at 6 hours, one wing
(from 124 hour old pupa) in dopa solution was well darkened
(deep gray) while the other wing in tyrosine solution had only
a light gray cast. This result would be expected because dopa is
an intermediate between tyrosine and melanin (oxidation of
tyrosine to dopa by tyrosinase is easily demonstrated — Evans
and Raper, 1937), dopa is more sensitive to enzymic oxidation
than many other chromogens (Schmalfuss, 1924), and dopa is
even readily oxidized nonenzymically, e.g. the 0.5% solution
begins to turn brown in a few days.
At 131 hours pupal age, dopa solution caused the whole wing
to become light gray in 3.5 hours with no selective darkening of
the future-black scales. At 160 hours pupal age, dopa solution
after 1.5 hours caused a wing to become gray with no differenti-
ation (except translucency ) of future-black scales; after rinsing
with water and air-drying overnight the wing was dark gray,
almost black, with markings barely discernible (Plate II, Figure
5). At about 185 hours pupal age, exposure to dopa solution
gave selective darkening that remained clear after the wing had
been rinsed with water and air-dried (Plate II, Figure 6). The
other (control) wing in deionized water did not darken (Plate
II, Figure 7). Figure 8 (Plate II) shows a dark scale from the
spot on a female wing (from 175 hour old pupa) which had
been artificially pigmented in dopa solution for 1.5 hours.
Minute spots of pigment are visible, seemingly within the sub-
8(3):69-90, 1969(1970)
WING MARKINGS
79
stance of the scale as claimed by Onslow ( 1916 ) , Reichelt ( 1925) ,
and Braun (1939).
When exposure to dopa solution was continued, for a pupa
about 190 hours old, the spot and apical scales were black
against a dark gray background at 24 hours, and at 48 hours the
wing was very dark gray, almost black, with markings barely
discernible (resembling Figure 5 n Plate II). The white scales
on the wing of a 204 hour old pupa, with markings recently
darkened, became very light gray after 4.5 hours in dopa
solution, and an even older pupa, apparently ready to eclose,
gave the same result.
The indication is that in earlier stages, e.g. 160 hours old or
less, all scales became pigmented at the same rate, while later
on (185 hours old or more) the future-black scales darkened
sooner but were eventually nearly equalled by the slower-dark-
ening future- white scales. Artificial pigmentation with dopa thus
is a “kinetic effect” resulting from the slower rate of darkening
of the future- white scales, not their inability to darken.
Other chromogens. — Cresols (54% m-, 29% p-, 17% other
phenols), DL-beta-phenylalanine, p-aminopehnol, resorcinol, and
catechol were tested as 0.5% solutions with the other wing from
each pupa (about 170 hours old) in 0.5% dopa solution. At 3
hours, all the wings in dopa solution were gray with future-black
scales darker gray. Phenylalanine and resorcinol gave no dark-
ening, the cresol mixture gave an orange tint to the basal half
of the wing but no darkening of future-black scales, p-amino-
phenol gave a tan-gray tint to the whole wing with doubt-
ful darkening of the markings ( translucency was difficult to
distinguish from pigmentation), and catechol gave an overall
orange-gray color with future-black scales darkened. These
results agree with the literature. The tyrosinase of the P. rapae
pupa oxidized catechol more readily than p-cresol (Pugh,
1934). Tyrosinase from the meal worm oxidized p-aminophenol
but not resorcinol (Gortner, 1910).
Inhibition by chemicals. Melanogenesis inhibitors (see Kol-
yer, 1966) were tested by adding at 0.5% to a 0.5% dopa solution,
with the other wing of each pupa (about 174 hours old) in 0.5%
dopa solution as a control (all turned gray with markings very
dark gray in 3 hours). Thiourea and L( + ) ascorbic acid allowed
no darkening of wing or markings, while the wing became light
gray but with little darkening of the markings with hydro-
quinone or 4-chlororesorcinol. Thiourea has been shown to
80
JOHN M. KOLYER
/. Res. Lepid.
cause pronounced inhibition of phenoloxidase activity in silk-
worm homogenates (Chmurzynska and Lech, 1963) and is a
well-known melanogenesis inhibitor. Ascorboic acid is a melano-
genesis inhibitor in vitro but is considered necessary in the diet
for optimum development of the silkworm (Ito, 1961). None
of these inhibitors prevented pigmentation when fed to larvae
in earlier work with P. rapae (Kolyer, 1966).
When the test was repeated (pupae about 177 hours old) with
the inhibitors (except hydroquinone ) at 0.05%, i.e. 10% on the
level of dopa instead of 100% as in the first test, the result was
partial inhibition (markings darkened but less intensely than in
the controls). Using pupae at about 193 hours old, at 0.005%
inhibitor (1% of dopa level) there was little, if any, inhibition.
Thiourea (at 0.5%) also inhibited darkening of the markings
(pupa about 193 hours old) in 0.5% catechol solution for 3 hours,
but the wing became pale orange-gray overall.
Inhibition by heat. — Gortner (1910, 1911b) reported that
activity of tyrosinase from the meal worm or the periodical
cicada is destroyed by heating at 75 °C for one minute, and
Onslow (1916) found that boiling the pupal wing of P. brassicae
prevented darkening in tyrosine solution. This denaturation of
the enzyme by heat was confimed for P. rapae as follows. One
wing from a 190 hour old pupa was placed in 1 ml water in a
small test tube, which then was immersed in water at 88-90°C
for 5 minutes. After 48 hours in dopa solution the wing was
pale yellow with no darkening of the future-black scales, while
the other ( unheated ) wing showed darkening of the future-black
scales in 2 hours.
In a series of hemolymph tests, the body fluid was squeezed
from three pupae (160 hours old) into a micro test tube, which
then was heated in water at 88-90° C for 5 minutes. The resulting
semisolid paste was diluted with 0.09 ml water and filtered by
drawing into a pipet plugged with cotton wool, and the filtrate
was applied to heated and unheated wings. Unheated, filtered
fluid similarly was applied to heated and unheated wings. At
48 hours, the unheated wing with unheated hemolymph was
pale tan with darkening at the torn base but no darkening of
the markings. The unheated wing with heated hemolymph was
gray-brown with darkening of the apex but not the spot. The
heated wing with unheated hemolymph was tan with markings
not darkened. The heated wing with heated hemolymph was
pale orange-yellow with no darkening. Though the markings
8(3):69-90, 1969(1970)
WING MARKINGS
81
in this series were not darkened when both hemolymph and
wing were unheated, as they were in other experiments, general
darkening of the wing was prevented only by heating both wing
and hemolymph. This is in agreement with the data of Onslow
(1916) and is explained by the presence of both oxidase and
chromogen in the wing as well as in the hemolymph.
Role of oxygen. — The hemolymph, which contains both oxi-
dase and chromogen, darkens when contacted with the atmo-
sphere (Pugh, 1934, and Wigglesworth, 1965: 383). In the
present work it was noted that darkening proceeded within a
minute when pupal hemolymph diluted with water was sparged
with fine bubbles of air.
In the enzymic oxidation of dopa, oxygen is necessary, though
only a trace is required (Gortner, 191 la, and Schmalfuss, 1924).
That dissolved oxygen was essential for the artificial pigmenta-
tions described above was indicated by soaking a wing (frorn
175 hour old pupa) in a 0.5% solution of dopa in essentially
oxygen-free deionized water (prepared by boiling 10 minutes
and cooling while bubbling in high-purity nitrogen). After 2
hours there was no darkening, even at the torn base, while the
other wing from the same pupa in ordinary ( air-containing ) 0.5%
dopa solution was blackened at the base and had the future-
black scales gray. The oxygen-free wing was rinsed with water
and allowed to dry in the air for 1.5 hours, during which time
artificial pigmentation proceeded, presumably due to availability
of oxygen (see Plate III, Figure 9).
3. Chemical Tests
Iodine solution. — Braun ( 1939 ) utilized a test ( treatment with
iodine solution, then zinc chloride solution) said to distinguish
between “hard” and “soft” chitin and found for Papilio ajax and
Ephestia kithnella that future-dark scales appeared light and
future-white scales appeared dark. This result, according to
Braun, “exhibits clearly that the different parts are found in
different stages of chitinization”. However, the test used by
Braun in not necessarily a specific test for chitin (Richards,
1947). Also, the hardest cuticles often contain less chitin than
the soft (Wigglesworth, 1965: 32). Richards (1947) treated
P. rapae scales with hot alkali and applied the chitosan test
(Campbell, 1929), which probably proves the presence of chitin
when it is positive, as it was for the dark scales. The white
scales were dissolved, but chitin in some cases is destroyed by
hot alkali, so that the presence of chitin in the white scales could
not be discounted.
82
JOHN M. KOLYER
/. Res. Lepid.
In the present work, pupal wings were treated with iodine
solution ( see Experimental ) as used in the chitosan test ( Camp-
bell, 1929). Chitin itself (Matheson Coleman and Bell practical
grade, prepared by purifying crab shells) was stained dark
brown by this reagent. For wings of pupae about 175 hours
old the spot was orange against a pale orange background after
iodine treatment, but translucency was a factor in this appear-
ance. Addition of 10% zinc chloride gave no color change, but
addition of 50% sulfuric acid caused rapid darkening of the wing
with, in some cases, marked resistance to darkening by the spot
so that it appeared as a 'window”. This may be the result of
absence of pterin pigments in the future-black scales, because
particles of leucopterin (from K & K Laboratories, Inc.) were
stained brown by treatment with the iodine solution when
followed by 50% sulfuric acid but not when followed by 10%
zinc chloride. Thus, pterin pigment distribution rather than
differences in chitinization possibly may explain the selective
staining reaction described.
Murexide test. — The forewings from a 189 hour old pupa were
exposed to chlorine gas for 9 hours and left in the air for 17
days. The wings, pink only around the edges, then were exposed
to gaseous NHs for 5 minutes, during which time they became
deep pink (rose color). The spots remained pale yellow against
the pink background (see Plate III, Figure 10), presumably
because the future-black scales were free of pterins such as
leucopterin, xanthopterin, and isoxanthopterin, which give the
murexide color (Ford, 1947, and Gates, 1947). Leucopterin,
isoxanthopterin, and other pterins have been found in the P.
brassicae pupa (Busnel and Drilhon, 1949, and Harmsen, 1966).
Ammoniacal silver nitrate. — The argentaffin reaction is used
to identify o-hydroxyphenols such as dopa (Richards, 1953),
which reduce the reagent to give free silver (black). Various
solutions have been described, all containing the readily-reduced
complex of silver ion with ammonia or amines. In the present
work Tollen’s reagent was found convenient.
Wings from 123, 125, and 131 hour old pupae were covered
with fresh Tollens reagent and within 5 minutes had turned
faint brown with no differentiation of the spot (which does not
become visible by translucency until about 135 hours). How-
ever, a wing from a female pupa about 175 hours old began to
darken immediately and in 2 minutes was practically black
with the two spots appearing as colorless "windows”. The other
8(3);69-90, 1969(1970)
WING MARKINGS
83
wing was soaked in water for 15 minutes before adding the
reagent; the result was the same except that the wing darkened
less (to brown rather than almost black). The wing from a
204 hour old pupa, with markings darkened, turned dark gray
after 2 minutes in the reagent so that the markings were barely
discernible. The same result was given by an adult wing (16
months old).
Selected white flakes of chitin were pale gray after 3 minutes
in the reagent, while particles of leucopterin turned brown to
black within 2 minutes. Urates, which are somewhat similar
chemically to the ammonium salt of leucopterin presumably
formed on adding Tollen’s reagent, are said to give a positive
argentaffin test (Richards, 1951: 71). Isoxanthopterin reduces
Tollen’s reagent (Gates, 1947).
Paper chromatography was done to verify the presence of
pterins in the pupal wing. Extracts were prepared from adult
wings (mixed sexes, washed with ether) or pupal forewings
(separate extracts for male and female wings from pupae about
190 hours old) by soaking the wings in 20% ammonia solution
for a few hours. The extracts were chromatographed vs. a
solution of leucopterin in 20% ammonia on Whatman No. 40 filter
paper by the ascending method (40 minutes at 78° F; solvent
front ran about 64 mm above point of application of extracts).
The solvent system was that of Partridge (1948) as recommend-
ed for pterins by Good and Johnson (1949). This was prepared
by shaking 40 ml n-butanol, 10 ml acetic acid, and 50 ml water,
allowing to stand 4 hours, and discarding the lower (aqueous)
layer. After drying, the paper was viewed under ultraviolet
light (mainly about 360 millimicrons). The adult wing extract
gave two fluorescent spots: very pale blue, R 0.11 (Good and
Johnson report 0.12 for leucopterin), and bright purple, R 0.28
(Watt and Bowden (1966) report 0.24 for isoxanthopterin).
These same two spots have been reported for adult wings of
Pieris rapae, P. brassicae and P. napi (L.). The pupal wings
also showed these two spots as well as an additional spot, pale
yellow, R 0.39, which was more intense in the female wing;
Good and Johnson assign this (R 0.38 ) to xanthopterin. Tollen’s
reagent poured over the paper caused intense darkening of the
spots of application of both the wing extract and the leucopterin,
indicating that much of the leucopterin applied to the paper
failed to migrate with the solvent.
The tentative conclusion is that the positive Tollen’s test is
caused by pterins and urates associated with the future-white
84
JOHN M. KOLYER
/. Res. Lepid.
scales. The weaker test after exposure of the wing to water
may have been due to partial extraction of pigment materials.
Formalin-sulfuric acid. — Le Rosen formalin reagent added to
wings from pupae about 190 or 198 hours old caused redwiolet
staining along the veins to a distance of about half way out on
the wing, which appeared clear pale-yellow. A fragment of
muscle tissue from the thorax showed red-violet streaks when
treated with the reagent. Crystals of dopa dropped in the
reagent also gave a red-violet color, as has been reported
(Deniges, 1926). Catechol gave the same result, but L-tyrosine
dissolved and then reappeared as a white precipitate, presum-
ably the sulfate, without giving a color. A red-violet color is
said to be given by phenols in general (Feigl, 1954). In the
present instance the eolor is attributed to phenolic substances,
such as dopa, in the hemolymph of the veins. No violet color in
the region of the markings was observed. An interesting inci-
dental effect was the clear display of the tracheae due to trans-
parency afforded by the reagent.
Coneentrated sulfuric acid, without formalin, gave no color,
nor was rapid dissolution of the seales observed as described
by Braun (1939). Incidentally, a technique for isolating the
wing membrane was provided by exposing the wing to the
acid for 5 minutes followed by a water rinse. It was then easy
to push away the scales as a soft mass.
4, Natural Pigmentation
Some experiments were performed to evaluate the suggestion
of Onslow (1916) that pigmentation is triggered by exposure
of the wing surface to air. When pupae with markings just
starting to darken, or even half-darkened, were placed in nitro-
gen, either dry or saturated with water vapor, pigmentation was
arrested. Results with isolated forewings were consistent. The
forewings were dissected from a female pupa with markings
judged just about to darken and placed in separate vials, one
filled with nitrogen and the other with air. A drop of water was
present in each case to saturate the gas and prevent desiceation.
After 7 hours the markings of the wing in air were about half
darkened, while no darkening had occurred in nitrogen. In a
similar experiment, male forewings, originally slightly darkened,
were left in humid air vs. nitrogen for 18 hours. The result was
further darkening in air vs. no change in nitrogen; see Plate III,
Figures 11 and 12.
8(3):69-90, 1969(1970)
WING MARKINGS
85
The pupal case was removed from one wing of a pupa with
markings half darkened. The imago eclosed 14 hours later with
markings fully darkened on the wing that had remained covered
but arrested at half darkened on the wing that had been ex-
posed. In a similar experiment, the markings were just starting
to darken when one wing was exposed. Again, the markings on
the exposed wing failed to darken, while the markings on the
covered wing were perhaps 75% darkened (in terms of final in-
tensity) after 4 hours.
Since the above results were attributed to desiccation by
evaporation of water from the exposed wing, pupae with the
apex of the wing exposed were placed in vials containing cotton
wool saturated with water to provide 100% relative humidity.
For air, results ( initiation of darkening, completion of darkening,
eclosion), in hours from start of experiment, were: 13, 20, 25;
1, 5, 15; 0, 5, 19. For oxygen, results were: 19, 24, 36; 0, 7, 13.
In all cases there was no difference at any time in appearance of
the exposed vs. the covered apex.
Using two pupae with markings not yet starting to darken,
the apex of one wing was exposed and covered with
petroleum jelly in an attempt to exclude air from the wing
surface. Approximate times (in hours as above) were: 31, 35, 38,
and 22, 27, 31. In the first case no difference was observed
during pigmentation of the apices, but in the second case there
was a delay in pigmentation of the outer part of the petrolatum-
covered apical marking. The fact that a delay was observed
suggests that contact of the scales with air is a requirement, but
the data of the preceding paragraph show that the apex can
be directly exposed to air for 13 hours (or to oxygen for 19
hours) before pigmentation commences.
CONCLUSION
Artificial pigmentation. — Oxidase must exist in both future-
black and future-white scales, since both eventually darken
when chromogen, e.g. dopa, is supplied. But why is artificial
pigmentation much more rapid for the future-black scales?
Braun (1939) argued that these scales are “softer” and “less
chitinized” but failed to prove chitinization of the future-white
scales. Even if selective chitinization were demonstrated, the
literature indicates this would not necessarily mean greater
hardness and lower permeability. Alternative possibilities to
greater permeability of the future- black scales are (1) less
oxidase in the future- white scales and/or (2) inhibition of
86
JOHN M. KOLYER
/. Res. Lepid.
melanogenesis in the future-white scales. The latter idea may
have merit on the basis that considerable evidence was given
by the chemical tests for the presence of pterins in the future-
white scales and the substantial lack of these pigments in the
future-black scales, and leucopterin, xanthopterin, and isoxan-
thopterin have been shown to have an inhibitory eflFect on potato
tyrosinase in vitro (Gonnard and Svinareff, 1951, and Isaka,
1952).
The 'relief stage” effect seems to indicate greater rigidity for
the future-white scales, but there is no reason to assign this to
"chitinization.” Hardening of the protein of the walls of the
scale, without chitin, seems as good a supposition since Richards
(1947) failed to demonstrate chitin in the white scales of the
adult P. rapae. It even seems possible that the white pigment
itself could have a reinforcing effect by being deposited in the
striations or corrugations of the scale.
In fact, the appearance of pigment at about 135 hours in the
future-white scales might explain all the observations. The
question then would revert to — What causes this selectivity
of deposition of pterin pigments?
Whatever the explanation, artificial pigmentation seems only
a relatively pale and less selective simulation of the natural
process in which some scales remain pure white while others
blacken intensely.
Natural pigmentation. — Braun states: "At a certain time in
development pigment is present in the body and the subsequent
dark parts, being still soft at this time, will deposit pigment”.
Presumably by “pigment” is meant chromogen. Also: “If at a
certain time in the development tyrosine is present, it will only
be deposited in those scales which represent a certain con-
dition of the chitin at this moment, which means only a certain
part of the pattern”. However, tyrosine is found freely in insect
blood (Brunet, 1963) and was found in the larvae and pupae
of Pieris brassicae (Stamm and Aguirre, 1955) and of the silk-
worm (Watanabe, 1956a, and Tomino, 1963 and 1965). Dopa
itself has been suggested as the chromogen in P. rapae ( Goodwin,
1965) and is present in all stages of the silkworm (Watanabe,
1956a and b). Thus, the chromogens tyrosine and dopa seem
to be present at all times. Furthermore, according to Buck
(1953), “There is reason to believe that enzyme, substrate, and
adequate oxygen are present together in the blood for some time
prior to the actual formation of pigment. The puzzle, therefore,
8(3):69-90, 1969(1970)
WING MARKINGS
87
is not so much in how melanin is formed, but in how its for-
mation in the blood of the intact animal is prevented and its
formation in cuticle so narrowly limited in time”.
The well-documented necessity for oxygen in natural pigmen-
tation was verified, and direct contact with the scales seems to
be required, which is consistent with the low capacity of hemo-
lymph to transport oxygen (Buck, 1953). However, Onslow’s
suggestion that pigmentation is triggered by air becoming
available due to pulling away of the wing from the pupal case
was discounted by removing a section of the pupal integument
at the apical region and finding delays of up to 13 hours in air or
19 hours in oxygen before pigmentation commenced. An inter-
esting example of oxygen supply as necessary but not sufficient
to initiate pigmentation is given by Fraenkel (1935) for the
newly eclosed blow-fly Calliphom erythrocephala. Pigmenta-
tion was inhibited and postponed by allowing the flies to dig
for an abnormally long time through sawdust in the presence of
air, showing that exposure to oxygen on emergence from the
pupal case was not sufficient to cause chromogen to oxidize but
that there is “certainly a nervous mechanism involved in initiation
of the coloraton process”. This nervous mechanism might func-
tion through a shift in oxidation-reduction potential of the
blood due to stress; see Buck, 1953.
In concluson, the complexity of the living system, both struc-
turally and chemically, makes dubious any simplistic mechan-
ism that might be proposed to explain pigmentation. Some
points can be demonstrated, but no general hypothesis, e.g.
that of Braun (1939), is very convincing when alternate ex-
planations can be suggested which also fit the limited data.
SUMMARY
Pupal wings of Pieris rapae (L.) were dissected and studied
at various times from pupation to eclosion (9-10 days). Scales
grew to full size from approximately 3 to 5 days with no ap-
parent difference between the future-white and future-black
varieties. At 5-6 days the wingcases became noticeably whitened.
During the next 3 days, before the onset of black pigmentation,
the presence of pterin pigments in the future-white scales, and
their substantial absence in the future-black scales, was indi-
cated by dark staining of the future-white scales with iodine
solution followed by 50% sulfuric acid, selective reduction of
ammoniacal silver nitrate by the future-white scales, and selec-
88
JOHN M. KOLYER
/. Res. Lepid.
tive pink coloration of the future-white scales by the murexide
test (chlorine treatment). Also, the future-black scales, lacking
pigment, were relatively translucent. A ‘relief stage”, as re-
ported by W. Braun (1939) for species including Papilio ajax
(L. ), was seen on brief drying of the wing in air; the future-
white scales appeared erect, the future-black scales collapsed.
Artificial pigmentation, reported by Braun using saturated tyro-
sine solution, was more effectively achieved with dopa. Another
successful chromogen was catechol. Before white pigmentation
at 5-6 days, all scales darkened in 0.5% DL-dopa solution at
the same rate, but in older pupae the future-black scales dark-
ened faster and so were blackened selectively at short times, e.g.
2 hours, though the whole wing became very dark by 48 hours.
This process is an enzymic oxidation requiring traces of oxygen
and prevented by melanogenesis inhibitors such as thiourea or
ascorbic acid or by brief heating of the wing at 90° C to destroy
the oxidase. Premature pigmentation also was achieved by soak-
ing the wing in pupal hemolymph, whereas darkening of the
future-black scales in water was at best faint. Thus, the data
indicate that the scales contain oxidase but are deficient in
chromogen. The reason for the pronounced difference in rate
of darkening in dopa solution between future-black and future-
white scales was not clear. Alternative explanations include less
oxidase in the future-white scales, greater permeability of the
future-black scales, and inhibition of melanin formation in the
future-white scales. The selective presence of pterin pigments
in the future-white scales possibly might explain not only
artificial pigmentation (pterins are known melanogenesis inhibi-
tors) but also the “relief stage” by reinforcing effect of pigment
deposited in the walls of the scale. The complexity of this
biological system, and the variety of explanations fitting the
limited data, make questionable the simplistic explanations that
have been proposed for black pigmentation in vivo, of which
artificial pigmentation is a pale and relatively nonselective
simulation. Pigmentation in vivo is not triggered by exposure
of the wing surface to air, as has been suggested (Onslow, 1916),
because darkening commenced and proceeded normally at times
of up to 19 hours after removal of pupal integument to expose
the apex of the wing to water-saturated air or oxygen. This
result, coupled with the observation of a delay in darkening
vs. the untreated apex when the exposed apex was covered
with petroleum jelly to exclude air, suggests that availability
8(3):69-90, 1969(1970)
WING MARKINGS
89
of oxygen at the wing surface is necessary but not suflRcient
to initiate the rapid (about 5 hours at 80° F) formation of
black pigment.
ACKNOWLEDGMENT
The author gratefully acknowledges the contribution of larvae
for this and other work by the Columbia, Missouri station of
the United States Department of Agriculture (Agricultural Re-
search Service, Entomology Research Division), where Mr.
Benjamin Puttier was Assistant Director and Mr. Richard K.
Morrison was in charge of the insectary rearing program.
LITERATURE CITED
ANONYMOUS. 1960. The Merck Index, 7th Ed. Merck and Go., Inc.,
Rahway, New Jersey.
BODENSTEIN, D. IQST Postembryonic development, in Insect Physiology,
edited by K. D. Roeder. John Wiley and Sons, Inc., New York Gity.
BRAUN, W. 1939. Gontribution to the study of the wing-pattern in Lepi-
doptera. Biol Bull 76(2): 226-240.
BRUNET, P. G, J. 1963. Tyrosine metabolism in insects. In: Pigment cell;
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JOHN M. KOLYER
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Journal of Research on the Lepidoptera
8(3):91-93, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
TWO NEW FORMS OF PLEBEJINAE
FROM WYOMING
CLIFFORD D. FERRIS
University of Wyoming, I^aramie
During the 1969 collecting season, two heretofore unde-
scribed forms of Plebejinae were taken. They are now briefly
described.
Plebejus ( Icaricia ) acmon lutzi form nov. psetidolupini
This insect is being described from a series of ten males. In
facies, the butterfly resembles L acmon lutzi dos Passos with the
exception of the submarginal orange spot-row on the secondaries.
Both ventrally and dorsally this row is broken into discrete small
spots resembling those on lupini (Boisduval). In several of the
specimens, the spots on the upper side are almost obsolete. The
black portion is clear, but the orange coloring is extremely re-
duced. Ventrally the orange spots are larger than those above,
but are much reduced over normal lutzi. The male genitalia are
identical to lutzi.
Holotype. — S , near Eagle Rock, 8200' approx., Sherman
Range, Medicine Bow N.F., Albany Co., Wyoming, 27 June,
1969.
Paratypes. — - 1 $ , 27 June; 2 S , 29 June; 2^,1 July; 4 $ ,
6 July, 1969. The paratypes are from Pole Mountain, 8200' ap-
prox., Sherman Range, Medicine Bow National Forest, Albany
Co., Wyoming. Expanse (costal margin length): 1.22 cm aver-
age.
Plebejus ( Plebejus ) saepiolus saepiolus
$ form nov. caerulescens
The description is based upon a series of seven specimens from
the type locality, which is in the Black Hills along the Weston
Co., Wyoming — Lawrence Co., South Dakota boundary. The
author has in his collection three additional females from Arizona
(Apache Co., vie. Alpine, 8200'-8500') which resemble the form
being described.
91
92
CLIFFORD D. FERRIS
/. Res, Lepid,
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8(3):91-93, 1969(1970)
NEW PLEBEJINAE
93
Two subspecies of saepiolus (Boisduval) are recorded from
the Rocky Mountain region: saepiolus (Boisduval) and whitmeri
Brown, although gertschi dos Bassos may intrude into the
western slope. “Normal” females of saepiolus are warm brown
dorsally with a submarginal orange spot-band on the secondaries
which varies from obsolete to relatively distinct. Occasionally
the ruddy female form rufescens ( Boisduval ) is taken. P. saepio-
lus whitmeri females show extensive dorsal blue of the same hue
as the males.
The new form represents a certain percentage of the female
population of s. saepiolus in the Black Hills. It diflFers from
normal females by having substantial dorsal blue scaling. On
the primaries this extends from the body over the basal half of
the wings; on the secondaries, the amount of blue varies in
extent and ranges from the basal half to the entire wing surface.
The dorsal submarginal orange spots on the secondaries may
or may not be present, as in usual saepiolus. Dorsally the brown
ground color is darker than in normal saepiolus, especially at
the apex of the forewing. The blue color generally suggests a
darker hue than that of the male and is quite luminous. It is a
deeper shade of blue than is found in whitmeri.
Holotype. — $ , Crooks Tower Road, Black Hills N. F., 6000'
approx., Lawrence Co., South Dakota, 4 July, 1969.
Paratypes. — - 3 $ , same date and location as holotype; 1 $
2 July, and 2 $ 4 July, 1969 from Weston Co., Wyoming — Law-
rence Co., South Dakota boundary along U. S. Highway 85.
Expanse (length of costal margin): 1.38 cm average.
The author would like to acknowledge a discussion with F.
Martin Brown which led to preparation of this paper. There has
been no distribution of paratype material to date, as the speci-
mens are needed for a continuing study of Wyoming Rhopal-
ocera.
This paper is published with the approval of the Director,
Wyoming Agricultural Experiment Station, as Journal Paper
No. 433.
Journal of Research on the Lepidoptera
8(3):94-98, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
CONCERNING THE NAMES AND STATUS
OF CERTAIN NORTH AMERICAN MEMBERS
OF THE GENUS PHYCIODES
J. W. TILDEN
125 Cedar Lane, San Jose, Calif.
In his “Synonymic List of the Nearctic Rhopalocera,” 1964,
dos Passos lists pulchella Bdv. (556c, p. 82) as a subspecies of
tharos Drury, with the more familiar pascoensis Wright as a
synonym. If the name pulchella actually applied to a western
entity, as Boisduval apparently believed it did, this listing would
be the valid one.
However, a study of all available references has convinced me
that the name pulchella cannot apply to any western population
of Fhyciodes tharos. The name pulchella first occurs ( Boisduval,
1852) as follows (my translation from the original French):
49. Melitaea pulchella
Pap. Tharos. Drury. Ins. I. pi. 21. f. 5.6.
It occurs in a large part of California. This species
should not be confused with tharos Cramer that also
(equally) inhabits the United States. It is well to note
also that morpheus Cramer, figured on plate 101, is
identical in every respect with that which was previous-
ly figured under the name tharos.
Since this is an indication that the figures cited depict what
Boisduval had in mind as pulchella, and since there is no other
description, the insect from which the figures were made may
be regarded as the type of pulchella Boisduval. Drury, in 1773,
could scarcely have had material from California. Edwards
(1864) states that Drury’s specimens of tharos, on which his
plates were based, came from New York. If this is true, no
figures of these New York specimens can form the basis of the
name of an entirely western population. It seems clear that
Boisduval’s name, pulchella, is a synonym of nominate tharos
Drury, and cannot apply to the insect we have known as pasco-
ensis Wright.
94
95
J. W. TILDEN
/. Res. Lepid.
It is difficult to know what Boisduval had in mind when he
proposed pulchella. No population of tharos is found over “a
large part” of California. If tharos occurs in California, it is
only in the northeastern corner of the state. The dark Phyciodes
of California is campestris Behr ( 1863 ) , at that time undescribed.
It is possible that Boisduval confused tharos and campestris, but
in no way did he suggest a name for what we now know as
campestris. It seems unlikely that Boisduval had specimens of
what he called pulchella, or he would not have needed to give
that name to a figure.
In 1869 Boisduval (Ann. Soc. Ent. Belg. 12:20, no. 50) men-
tions Melitaea pulchella again:
50. Melitaea pulchella, Boisd.
Papilio Tharos, Drury, Ins. 1. PI. 21, f. 5-6.
Well scattered (assez repandue, or distributed) in cen-
tral California. This species should not be confused
with Tharos Cramer which inhabits certain parts of
North America.
And again in 1869, Boisduval (ibid. 12:53, no. 37) writes of
Melitaea tharos Boisd. et Leconte (!?), gives Argynnis tharossa
Godt., as a synonym, and again says of tharos that it “occurs
also in certain localities in California.”
And finally, in the same work, next number (no. 38) he lists
Melitaea cocyta Cramer (now considered a synonym of tharos)
with Argynnis morphea Godt. as a synonym. Of morphea he
says, “It was captured at Los Angeles.”
These references indicate that Boisduval persisted in thinking
(a) that pulchella was different than tharos, and (b) that both
tharos and pulchella occurred in California,
It is interesting to note that neither of the common lowland
California species of Phyciodes (mylitta and campestris) were
among the material sent to Boisduval by Lorquin and described
by Boisduval in 1852. This strengthens the inference that the
earlier Lorquin collections were made in the mining country of
the Sierra Nevada, rather than in the Bay Region of California.
It is suggested that caution be used in fixing San Francisco as
the type locality of species described by Boisduval in 1852.
In his Synonymic List ( 1964 ) dos Passos listed mata Reakirt
as a subspecies of mylitta Edwards, with harnesi Skinner as a
synonym, but more recently (Journ. Lepid. Soc., 23:120) he
places mata as an aberration of P. campestris Camillas Edwards
(569b). The checkered history of this name, given to a very
unusual appearing single specimen, is interesting. Reakirt de-
scribed it as a bleached specimen which nevertheless he con-
sidered to represent a distinct species (Reakirt, 1866). Strecker,
(1874) says of this type of mata, “Female. Expands lYz inches.”
8(3):94~98, 1969(1970) AMERICAN PH'^^COIDES
96
Brown (1966) devotes an illuminating paragraph to the mata-
pallida problem. He considers the type of mata to be albinic
rather than faded (an opinion expressed earlier, by Strecker).
Brown states, “if it is mylitta, it is unusually small.” Fropi this
I judge that Strecker s measurement of “V/z inches” is very ap-
proximate, since this is very large for a mylitta.
Brown (loc. cit. ) finds it impossible to decide whether mata
belongs to the concept of mylitta^ or to camillus. This seems
to have been the reaction of all who have discussed this speci-
men. Reakirt thought it faded; Strecker and Brown thought it
not faded; Barnes & McDunnough (1916) thought it to be
mylitta, both worn and faded when taken. None seems to
agree. The recent action by dos Bassos disposes of the name as
populational. This seems far better than to use the name mata
to affect other better established names.
Concerning the status of the names pallida Edwards and
barnesi Skinner, which have traditionally been associated with
mylitta Edwards, there is what appears to be good biological
and distributional evidence that mylitta and pallida are dis-
tinct species, with barnesi a weakly differentiated subspecies of
pallida. Here is the evidence: pallida and barnesi are one-
brooded. Mylitta is holodynamic wherever found, breeding con-
tinuously as long as weather conditions permit. In Utah and
northwest into Washington, both one-brooded populations
{pallida-barnesi) and multi-brooded populations (mylitta) are
sympatric and separable when once known by subtle markings
as well as by size. The pallida-barnesi complex are consistently
larger insects, and have a dark spot in cell Cus of the forewings
that shows on both upper and lower surfaces, in most specimens.
In addition, the females of pallida-barnesi show a more or less
complete row of outer crescents on the underside of the hind
wings, these crescents creamy or buffy, and no one of them
much darker than the others.
F. mylitta averages smaller, is multi-brooded over its entire
range, lacks the dark Cuo spot in most specimens and the fe-
males, as in the males, have one of the crescents on the under-
side much darker than the others, the typical “crescent spot.”
Populations of pallida-barnesi and of mylitta, when sympatric,
are not synchronic. The single brood of the pallida complex
peaks at a different time than any of the several broods of
mylitta.
These pieces of evidence convince me that mylitta Edwards
1861 should be considered one species, and that pallida Edwards
1864 should be regarded as a separate species, with barnesi
Skinner 1897 as a western subspecies of pallida.
97
J. W. TILDEN
/. Res. Lepid.
The type locality of pallida Edwards was fixed by Brown
(1966) as Flagstaff Mountain, Boulder Co., Colorado. The
stated type locality of barnesi Skinner is Glenwood Springs, Gar-
field Co., Colorado, far west of the Continental Divide and cli-
matically allied to Utah. F. pallida barnesi extends south from
the type locality to northern Arizona and northwesterly to Wash-
ington and southern British Columbia, east of the Cascades.
Over much of this range it occurs with mijlitta. I have examined
sympatric material of these species. Lack of similar material
from the higher eastern parts of Colorado suggests that true
mylitta either does not extend there, or is rare there, or that the
distinctions between mylitta and pallida may have been over-
looked. I favor the first hypothesis. Plentiful material that I
have examined from eastern Colorado seem to me to be all
pallida. Genitalic distinctions are either minor or nearly lacking
between these two species but may be demonstrated by further
studies.
Changes in the listings of our Phyciodes have been frequent
but the following seem justified:
566. tharos (Drury) 1773
a. t. tharos (Drury) 1773
pulchella (Boisduval) 1852
(return to former synonymy)
b. t. arctica dos Passos 1935
c. t. pascoensis Wright 19p5
and: 571.1 pallida (Edwards) 1864
a. p. pallida (Edwards) 1884
b. p. barnesi Skinner 1897
572. mylitta (Edwards) 1861
The status of any populations that may belong under mylitta
does not form a part of this paper, but will be treated separately
by Mr. David Bauer.
I am grateful to Mr. David Bauer for critical review of the
manuscript and for many valuable suggestions. He has been
kind enough to allow me to read the manuscript of a forthcoming
paper in which he expresses the same conclusion regarding the
specific status of Phyciodes pallida (Edwards).
REFERENCES
BARNES, WM., & F. BENJAMIN. 1926. Bull So. Calif. Acad. Sci,
25( 1 ) :13, under no/ 251.
BARNES, WM. & JAMES McDUNNOUGH. 1916. Contrib. Nat. Hist.
Lepid. N. A., 3(2):93^97.
1917. Check List Lepid. Bor. Amer., p, 10, under no. 249.
BOISDUVAL, JEAN. 1852. Ann. Soc. Ent. France (2)10(2) : 306, no. 49.
1869. Ann. Soc. Ent. Belg., 12:20, no. 50.
BROWN, F. M. 1966. Tram. Am. Ent. Soc. vol. 92:443-448.
8(3):94-98, 1969(1970) AMERICAN PHYCOIDES
98
DOS PASSOS, CYRIL F. 1962. Journ. Lepid. Soc. 15(4) :219.
1964. Mem. Lepid. Soc. I, p. 82, no. 566c; no. 559b; and no. 559c.
1969. Journ. Leiiid. Soc. 23(2) :115-125.
DRURY, DRU. 1773. Illust. Nat. Hist., pi. 21, ff. 5, 6.
DYAR, H. G. 1902. Bull. U. S. Nat. Mus. No. 52, p. 20, under 191.
EDWARDS, W. H. 1864. Proc. Ent. Soc. Phila., 2:505.
1869. Trans. Am. Ent. Soc. 2:207-209.
1872. Synopsis N. Am. Butt., p. 17.
1877. Trans. Am. Ent. Soc., 6:26.
1884. Trans. Am. Ent. Soc. 11:192, under no. 192.
HOLLAND, W. J. 1931. Butt. Book, rev. ed., p. 135, under no. 1.
KIRBY, WM. F. 1871. Syn. Cat. Diurnal Lepid., p. 172, under no. 12.
McDUNNOUGH, JAMES. 1938. Mem. S. Calif. Acad. Sci. I, p, 19, under
no. 265.
REAKIRT, TRYON. 1865. Proc. Ent. Soc. Phila., 5:226-227.
1866. Proc. Ent. Soc. Phila., 6:142.
SCUDDER, S. 1875. Bull. Buff. Soc. Nat. Hist., 2:266, no. 166.
SKINNER, H. 1898. Syn. Cat. No. Amer. Bhopal., p. 17, under no. 117.
1897. Can. Ent. 19:155.
STRECKER, H. 1874. Lepid. Bhopal. & Heter., (8):65-66, pi. 8, fig. 11.
1878. Butt. & Moths of N. Amer., p. 121, no. 233b and no. 133.
1900. Lepid. Bhopal. & Heter., SuppL, (3):23.
WRIGHT, W. G. 1905. Butt. West Coast, p. 165, pi. XXI, ff. 198, 198a.
Journal of Research on the Lepidoptera
83):99-104, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S,A, 91006
© Copyright 1969
THE LIFE HISTORY OF
AMBLYSCIRTES LINDA ( HESPERHDAE ) ‘
J. RICHARD HEITZMAN^ and ROGER L. HEITZMAN
3112 Harris Avenue, Independence, Mo.
Five forest dwelling species of Amblyscirtes are known to
occur in the Missouri-Arkansas plateau region. Of the five, Am-
blyscirtes linda H. A. Freeman is the most eremitical. We have
found this species a habitue of undisturbed woodland areas
along or near small streams with abundant colonies of the host
plant. Although the foodplant, Uniola latifolia Michx., occurs
in most of the Midwestern and Eastern states, the northern range
of A. linda seems to be the southern tier of counties in Missouri.
The Gramineae host species is one of the favorite lepidoptera
foodplants of the region acting as primary or secondary host for
Amblyscirtes belli H. A. Freeman, Amblyscirtes samoset Scudder,
Amblyscirtes vialis Edwards, and Lethe anthedon Clark. A. linda
females can occasionally be collected from low blooming flowers
of the wild blackberry and are easily induced to oviposit in cap-
tivity. The progeny of early May females have invariably de-
veloped rapidly with a 100% emergence of the imagines in June.
We have never encountered a third brood in the field and if one
occurs it is probably vestigial as with A. belli. The adult larvae
are unique in several aspects from the other members of the
genus that we have studied. The freshly emerged imagine is
distinctive with bright fulvous scaling on the dorsal and ventral
surfaces of the forewing. Worn specimens lose this fulvous scal-
ing and are then easily confused with A. vialis and A. belli.
The following description is based on studies of over 100
reared and field collected larvae from northern Arkansas and
southern Missouri localities. The illustrations were made by the
junior author from specimens reared from a female taken 3 May,
1964 near Eureka Springs, Arkansas.
^Contribution No. 167, Entomology Section, Div. of Plant Industry, Florida Depart-
ment of Agriculture and Consumer Services, Gainesville.
2Research Associate, Florida State Collection of Arthropods, Div. of Plant Industry,
Florida Department of Agriculture and Consumer Services.
99
100
HEITZMAN and HEITZMAN
J. Res. Lepid.
Figs. 1-6. — Amblyscirtes linda H, A. Freeman. 1, setae of first instar sur-
anal plate; 2, first instar setal maps of prothorax, mesothorax, eighth ab-
dominal segment, all in left lateral aspect; 3, first instar head capsule,
frontal aspect; 4-5, mature larva head capsule, frontal and left lateral aspect
showing position of stemmata; 6, ovum X 25.
8(3):99-104, 1969(1970) AMBLYSCIRTES LINDA
101
OVUM: Width .90 mm, Height .65 mm, As pictured, unmarked
shiny white in color. Ova are laid singly on the under surface of
a leaf near the edge. The egg shell is devoured upon emergence.
FIRST INSTAR LARVA: Head and prothoracic shield shiny
black. Integument white with minute white setae. After de-
vouring the egg shell the larva moves to the edge of a leaf and
makes a small tent shelter by folding the edge of the leaf par-
tially over and fastening it with strong silken strands.
SECOND INSTAR LARVA: Head and prothoracic shield shiny
black. Body color pale bluish green covered with minute white
setae. There is a faint indication of a middorsal line. Larval
tent as in first instar but longer, in some cases as much as 30 mm.
in length.
THIRD INSTAR LARVA: Head white with dark brown lines
circling the edges of the epicranial plates and covering the pos-
terior region of the head. Labrum and mandibles brown, cly-
peus white. Body color pale bluish white with a thick covering
of microscopic white setae. Prothorax paler with a constrastingly
shiny black prothoracic shield. Thoracic spiracle black, abdomi-
nal spiracles inconspicuous. There is a pale blue middorsal heart
line fading posteriorly. An entire leaf is used for the tent in this
instar which is folded in half and sealed along the edges. The
tent is then devoured from the tip down.
FOURTH INSTAR LARVA: Head white, mandibles and labrum
reddish brown, clypeus white with a fine black center line. Mid-
cranial inflection black edged, wider along the laterofacial su-
ture. A dorsally directed black dash extends parallel to the wide
central band on each side rising from the laterofacial suture
band and stopping a few millimeters short of the vertex. Pos-
terior region of head black ringed. Another wide black band
circles the edges of the head beginning ventrally just below the
anterior stemmata then rising dorsad to the vertex, narrowing
where intersected by the midcranial inflection. Head finely
setose. Body gound color bluish white, entirely covered dorsally
by very short white setae. The posterior end of the abdomen has
longer pure white setae. There is a blue middorsal line fading
posteriorly and a faint white stigmatal line. Thoracic spiracle
black, abdominal spiracles small, white ringed. Abdominal area
pale bluish grey. Prothorax pure white with a thin black pro-
thoracic shield. Larval tent as in the third instar.
FIFTH INSTAR LARVA: Length of mature larvae 25 to 30 mm.
Width of head case 3.5 mm. Body ground color pale bluish
white so thickly covered with snow white setae that the entire
body appears covered with snowy mold. There is a faint blue
102
HEITZMAN and HEITZMAN
/. Res. Lepid.
Fig. 7-9. — Amblyscirtes linda H. A. Freeman. 7, adult male
ventral and dorsal view of specimens from Eureka Springs,
mature larva in opened larval tent, natural size; 9, pupa,
right lateral aspect X 4.
and female,
Arkansas; 8,
ventral and
8(3):99~104, 1969(1970) AMBLYSCIRTES LINDA
103
middorsal line that disappears posteriorly. Abdominal area
slightly bluer with fewer, shorter setae, Spiracles as in fourth
instar. Thoracic legs very pale orange brown. Prothorax white
with prothoracic shield pale grayish white with two narrow
dark subdorsal marks. Head white, mandibles reddish brown,
clypeus white with vertical line and lateral bordering sclerites
black. Head banded on each side with a wide black line, ven-
trally enclosing the four anterior stemmata then running dorsad
across vertex where it narrows. Midcranial inflection widely
banded with black as are the laterofacial sutures. A pointed
black band rises from each of the laterofacial suture bands. The
paraclypeal spines ( Klots 1966 ) are well developed, arising from
a position ventro-lateral to the angle of the clypeus and angled
ventrad. Of the four anterior stemmata, 3 is the largest, 4
slightly smaller, 2 is slightly smaller than 1 but protrudes twice
as far, 6 is almost directly caudad of 3, 5 is ventrad of 6. At
maturity the head markings are partially obliterated by an ex-
tremely thick covering of short white setae.
The final instar larval tent is composed of an entire leaf folded
over and sealed along the edges with silken strands. Both upper
and lower ends are left open. Adjacent leaves as well as the
tent leaf are entirely devoured. Larvae are often observed feed-
ing openly in the daytime, especially in native woodland habi-
tats. The larvae are unusually docile, showing no agitation when
touched or handled.
PUPAL SHELTER: When the larva is ready to pupate a fresh
leaf is rolled over and sealed along the edge and both ends.
The larva spins only a vestigial silk lining. The tent is fastened
at the upper end to a grass stem two or three inches above the
ground. No instances of the shelters being allowed to fall to
the ground have been observed although this has been the
accepted practice with other Amhlyscirtes species we have
reared. Pupation occurs three to four days after shelter construc-
tion.
PUPA: Length 17-19 mm., width at wing cases 4.5 mm. Color
of wing cases bright creamy yellow, abdomen paler with a whit-
ish cast. Head and eye cases slightly darker with many stiff
reddish setae, a few of which extend over onto the thorax.
Mesothoracic spiracles bright red and conspicuous. The abdom-
inal segments have a sparse covering of short orange-red setae
arranged in definite tufts on each segment paralleling the tounge
case which is pale orange brown, long, and slightly curved.
Cremaster reddish brown, curved ventrally with several long
stiff bristles. The cremaster hooks are firmly embedded in the
104
HEITZMAN and HEITZMAN
/. Res. Lepid.
side of the shelter and hold the pupa in fixed position at the base
of the tent.
Our special thanks are due Dr. Alexander B. Klots, American
Museum of Natural History, and Dr. Howard V. Weems Jr.,
Florida State Department of Agriculture, for reading the manu-
script and making helpful suggestions.
LITERATURE CITED
KLOTS, ALEXANDER B. 1966. The Larva of Amhlyscirtes samoset
(Scudder) ( Lepidoptera: Hesperiidae). Jour. New York Ent. Society,
Vol 74, No. 4:185-188.
Journal of Research on the Lepidoptera
8(3):105»117, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
FURTHER OBSERVATIONS ON
“HILLTOPPING” IN PAPILIO ZELICAON
RICHARD GUPPY
Thetis Island, British Columbia, Canada
In the past i have wbitten two papers in which I set forth
the “mating rendezvous” theory to account for the hilltopping
habits of certain butterflies. I3uring the ensuing years many
things have come to my notice that have sapped my confidence
in this theory. After coming to live on Thetis Island in 1965, I
found myself in an excellent position to- study the habits of
Papilio zeiicaon, which is by far the most notable and persistent
hilltopper. The results of my observations here have caused me
to abandon the mating rendezvous theory entirely.
Later, in correspondence with Mr. Oakley Shields of San
Diego,' California, I learned that he was making a detailed study
of P. zelicaon, with a view to publishing a paper on the hilltop-
ping habits of this species. I felt that, owing to my limited op-
portunities for research, I could do better by collaborating with
Shields, rather than writing up my own findings. He accepted
my suggestions for this plan, and for over a year I sent him
several long letters detailing my observations. In his paper,
“Hilltopping” (196), he referred the reader to my published
papers which I had repudiated.
I conceived the idea of crudely marking, and releasing, a few
male F. zelicaon, I first considered this as a preliminary test,
intended mainly to show whether the recovery ratio would be
sufficient to warrant a more elaborate program. However, I
now feel that the results achieved are worthy of publication.
Thetis Island consists of two ridges running approximately
north, and south, each about 4 or 5 miles long and reaching 600'
at the highest point. Between these ridges the valley is for a
large part below, or barely above, high tide level. In the south-
ern part of this valley an area of about 10 acres is almost flat,
105
106
RICHARD GUPPY
/. Res. Lepid.
supporting a rank growth of swamp loving herbage, mainly
sedges (Carex spp.) with a considerable admixture of the
water parsley {Oenanthe sarmentosa) . Undoubtedly, in this
swamp most of the P. zelicaon population of island feed as
larvae.
The land rises quite evenly from the swamp to the summit of
the west ridge (Birchall Hill). This hill is close to, and easily
seen from, the swamp, and so should provide the most likely
place to find hilltopping butterflies. In fact, it is heavily wooded
and I have never seen any butterflies there. From the east
side of the swamp rises a sheer cliff 100' high, blocking off all
view of the east ridge (Moore Hill). Between this cliff and
Moore Hill the land is irregular with many small ridges and
valleys. It seems improbable that any insect could find the hill
by following land contours, as suggested in my paper on Oeneis.
Moore Hill is very sparsely wooded at the extreme summit, and
on the gradual slope extending northward. It is much frequented
by butterflies, chiefly Papilio zelicaon, P. rutulus, and P. euryme-
don.
My system was to clip off the tip of one forewing of butterflies
to be released. Those collected on Moore Hill had the left wing
clipped, those collected in the swamp the right wing. As an
additional check, I clipped the tail from the hind wing opposite
to the clipped forewing. Due to the frequency with which
Papilios lose the tails by accident, I did not plan to draw any
conclusions from insects with missing tails, unless this mark was
clearly supported by a clipped forewing. In any event I caught
only one individual which had lost the tip of a forewing, other
than those which were clearly my released specimens. This one
had part of a wing removed leaving a ragged edge, not neatly
clipped as in my marked specimens. Both tails were intact.
All others, counted as recaptures in the ensuing account, had
neatly clipped forewings, and the opposite tail only missing.
In addition to my marking and releasing program, I kept
records to ascertain the average time needed to collect, respec-
tively, a male and a female P. zelicaon, on the hilltop and in the
swamp. I also tried to estimate the number of P. zelicaon usually
present on the hilltop; as there were very few, this was not
difficult. Due to the large area and the large number of butter-
flies present, no estimate that would be of any value could be
made of P. zelicaon numbers in the swamp. On the hill I noted,
each day that I collected there, (a) the largest number of P.
zelicaon seen together at one time, and ( b ) the minimum number
8(3): 105-1 17, 1969(1970)
HILLTOPPING
107
that could have visited the area during my stay. The last figure
was arrived at by adding to the number collected any others
seen after I had ceased collecting.
The areas which I collect at a given time are decided mainly
by mercenary considerations. When P. rutulus and P. eurymedon
are present I collect the hill; when only P. zelicaon are available
I collect the swamp. As P. zelicaon is by a week or two the
earliest on the wing of the three species, I commence the season
by collecting the swamp, but soon resort to the hill until all
species become too worn to be worth any further effort. When
the late brood of P. zelicaon emerges, the other two species
being single brooded, I collect the swamp for the remainder of
the season. Thus I did not manage to check the swamp for the
possible return there of specimens taken on the hill.
All releases were carried out in my garden. Unfortunately,
I do not have any map of a scale that would allow of my calcu-
lating exactly the distances involved. The distance to the swamp
is a little over a mile; the distance to the hilltop several times
farther, a circumstance which can hardly account for the results
as follows. The first P. zelicaon of the season I collected in the
swamp on May 8; these were 2 females, not absolutely fresh.
From that date up to May 18, I alternated between the hill and
the swamp. The first release from the swamp I made on May
17, but as I did not return to the swamp until June 5, later dis-
closures will show that I had little opportunity to recapture this
one. On June 5, I released two from the swamp, but returned
there only one more day, June 7. On that day, I released three
from the swamp.
From May 14 to May 25, I was at the hill on alternate days.
I released one butterfiy on each of the following days: May 16,
18, and 21, On May 25, I released two. On May 18, I recap-
tured a marked specimen in my garden. I had not yet made
that day’s release, so this specimen was the one released two
days earlier; it had made no attempt to travel anywhere. Fol-
lowing the May 25 releases, I did not get back to the hill for
five days. I was then at the hill May 30, June 2, 4, 8, 10, 11,
and 13. I released one specimen June 2 and one June 8. On
June 10, I recovered one specimen which had been marked and
released from the hill; this was the only one retaken while col-
lecting on the hill. I took this one home again, clipped the other
wing, and released it; on the two subsequent days I returned to
the hill I did not see it. Early flight P. zelicaon were now mostly
108
RICHARD GUPPY
/. Res. Lepid.
worn, so I terminated my spring collecting on the hill.
On July 13 I noted second brood P. zelicaon on the wing and
therefore tried the swamp again. On that day I released four
butterflies. I did not get back to the swamp again until July 16,
when I recovered one of the July 13 releases. I took this one
home, clipped the other wing and released it again, together
with the three others captured on the same day (July 16). On
my next visit to the swamp, July 18, 1 recovered two once
marked specimens. Both of these, I double marked and released
again. On my next and last visit to the swamp, July 21, I re-
covered one twice marked specimen. On that day, 1 also
recaptured a twice marked specimen at my garden.
In summing up, I think I may be permitted to ignore the six
spring brood specimens from the swamp which I released.
I made almost no effort to recover these; at that time I supposed
that if I did see them again it would be on the hill. Thus we
have a recovery rate of one out of seven released from the hill,
this one being double marked and not seen again. From the
swamp, we have a recovery of three out of seven, these three
being double marked and one of them recovered again. In
view of the small number of butterflies used, and the much
greater distance from the release point to the hill, I can hardly
claim that these figures offer convincing evidence to the effect
that the butterflies returned more readily to the swamp than
to the hill. Still, I think that they can be regarded as suggestive.
In any case, this is irrelevant to the theory that I set out to
gain evidence on, namely, that butterflies from the swamp had
no interest in the hill. I had as much opportunity of recovering
the six early releases from the swamp, had they gone to the
hill, as I had of recovering the seven released in the summer.
My data for collecting success are also interesting. They
show that in the swamp I required 15 minutes to collect a male,
and 50 minutes to collect a female. On the hill I required 28
minutes to collect a male, and 5 hours 45 minutes to collect a
female. Actually these figures are highly misleading in favor
of the hill. On the hill the butterflies are concentrated into
about half an acre of level or moderately undulating ground,
with a few scattered trees, and only very short grass or mosses
between them. The butterflies pass repeatedly over a largely
predictable course and so are easily intercepted. In the swamp
they are scattered over 10 acres, mostly covered with knee-high
saw-edged sedges. This area was at one time cultivated as a
market garden, and to keep it drained, it was crisscrossed with
8(3):105-117, 1969(1970)
HILLTOPPING
109
a system of ditches up to two feet deep. Through many years
of neglect these ditches have ceased to provide drainage, but
many are still effective as traps for unwary butterfly collectors.
There is no hope of running through this mess, and no sure
way of predicting the course that any butterfly will take, in
order to intercept it. Of course, there is no basis on which I can
estimate accurately the effect of these handicaps on my col-
lecting success, but as a guess I should say that the opportunity
of securing any particular butterfly is ten times as great on the
hill. From my records I find that in fact I never saw more than
two P. zelicaon together on the hill, and it is possible that no
more than six in all were there during any of my visits lasting
from one to three hours.
It will be realized that to make this collecting at all profitable,
I was taking mostly other species, and so it may be supposed
that my poor success with P. zelicaon was due to such distrac-
tions. Actually, since P. zelicaon are much easier to net than
P. eurymedon, and much more in demand among collectors than
P. rutulus, I always concentrate on any P. zelicaon that show up,
going for the others when no P. zelicaon are in sight. Again,
after my elaboration of the difficulties of collecting in the
swamp, it may be wondered how I was able to recover so many
marked specimens from among the large number scattered over
the area. I think the explanation here is that the insects re-
turned, not just to the swamp, but to a particular small area that
they had staked out as territory. They may travel over a con-
siderable distance, but as a sort of patrol, continually passing
and repassing over the same track. There are in the swamp
some patches of slightly higher ground that are dry, and lack the
heavy cover of sedges. I soon learned that I attained as much
success by staying on these dry patches and trying to intercept
any passing insects, as by running all through the sedges and
falling into the ditches. It can be seen that if a butterfly re-
turned to its regular patrol, I would collect it again.
There is one evident conclusion to be drawn from these
observations. The hilltop is not a permanent attraction for a
large part of the P. zelicaon population. It is just a good col-
lecting place because it is a very small area in which the pres-
ence of a few butterflies can be reliably predicted. When, as
is the case here with P. eurymedon and P. rutulus, the host
plants are scattered thinly over a wide area, the hill may be
the most productive collecting site. But when, as with P. zelica-
on here, a large proportion of the available host plants are con-
no
RICHARD GUPPY
/. Res. Lepid.
centrated into a relatively small area, this emergence area
provides, on the whole, a better collecting site than does the hill.
The small number of males on the hill could be explained
by postulating that these are dominant or successful individ-
uals, which drive all others off. But it is not so easy to account
for the almost complete absence of females. In fact, there is
no evidence that the hill has any attraction at all for females.
I am sure that if I were to mark out half an acre of grassy land
anywhere on Thetis Island, and spent five hours there, during
the flight season of P. zelicaon and while temperatures were
favorable to butterfly activity, I could not fail to collect one
or more females. In my garden, where there are a fair number
of flowers attractive to butterflies, I would quite certainly do
much better than that.
When expounding the mating rendezvous theory in my Oeneis
nevademis paper, I supposed, with some justification, that O.
nevadensis was a rather rare insect in the area of my observa-
tions. This, added to the fact that females would leave the
hilltop immediately after copulation was ended, nicely ac-
counted for my seldom collecting any there. But for a species
as common as P. zelicaon is on Thetis Island, this theory will
not do at all. Every female would have to make at least one
visit to the hilltop. Shields, in a letter, has suggested that I do
not collect the hill at the right time of day. I have frequently
been there in the morning when butterflies were barely starting
to move. On the British Columbia coast, where spring nights
are always cool, this does not by any means require early rising.
There remains the late afternoon. But it is obviously impossible
for the insects to predict their time of arrival at the hilltop.
One cannot imagine a whole flock of females hiding just down
the slope somewhere, waiting to pop up at a given signal. If
the collector remains on the spot until mid-afternoon, as I have
often done, and no females have shown up, it is safe to predict
that there will be very few there that day.
My mating rendezvous theory as set forth in my earlier
essays depended on the proposition that the butterflies con-
cerned emerged from the pupae as a few individuals widely
scattered. To P. zelicaon on Thetis Island, this cannot apply.
Plenty of both sexes can be seen at the swamp, and it is quite
evident that none of them are headed anywhere in particular.
The females are ovipositing and the males are looking for
females. I have observed many courtship flights, but seen few
actually in copulation. My failure to observe actual pairs may
8(3): 105-1 17, 1969(1970)
HILLTOPPING
111
be largely due to the fact that I would sooner collect the insects
than wait to see what they are going to do. But it is probable
that most butterfly courtships end abortively simply because
females are receptive only for short periods. Whatever the reason
for the scarcity of copulating pairs, it obviously cannot be be-
cause the females are staving off their suitors until they can get
to the hilltop. The advantage accrueing to those that “cheated’"
would be tremendous. They would avoid waste of time and
effort entailed in a long hazardous return journey. Shields’ idea
that there is an advantage in stabilizing the gene pool is not
very convincing. This makes it one of those cases where a habit
not beneficial to the individual becomes established because it
is of benefit to the population as a whole. I do not wish to
wander off here into a long discussion of this concept. It must
suffice to say that such a habit must be neutral or at the worst
only slightly detrimental in its effect on the individual, other-
wise it could not persist long enough to become established in
the population.
It would be foolish, of course, to claim that Shields’ experi-
ment proved nothing at all. The fact that his butterflies some-
times returned to hills other than those from which they were
taken, shows that the homing instinct is not entirely responsible
for his recovery of marked specimens. His theory of hilltopping
by direct view of the hill is far better than my idea of insects
following the ground contours, as set forth in my Oeneis paper.
But it forces the conclusion that insects cannot reach a hilltop
until they come by accident to a point from which they can
see it. It could hardly be of much benefit to males to spend their
time waiting on a hilltop for females, a large proportion of
which would never get there. Shields, as his illustration plainly
shows, was able to work on neat little humps sticking out of a
nearly level and largely treeless plain. He would almost cer-
tainly have obtained different results if he had met with such
a situation as pertains here, where very few summits can be
seen before you are almost up to them, unless from certain
points of vantage. A reasonable supposition would be that the
hilltopping instinct becomes dulled under the latter conditions.
But could this circumstance almost entirely eradicate an in-
stinct which was of any great advantage to the possessors?
There is a definite relationship between the number of P.
zelicaon commonly on a given hilltop, and the availability of
food plants. On the hill at Wellington, which was much used
by P. rutulus, P. eurymedon, and Oeneis nevadensis, I saw no
112
RICHARD GUPPY
J. Res. Lepid.
more than a dozen P. zelicaon in nearly 20 years that I collected
there. Yet they were not entirely absent from the surrounding
country, and I often found a few larvae on parsley in my
garden. Mt. Benson, a very conspicuous lone summit, on most
of my visits showed only two or three P. zelicaon at the summit.
But on Mt. Prevost, which offers similar attractions, there are
seldom less than fifteen or twenty. It is true that I have not
discovered the source of this comparatively large population of
F. zelicaon on Mt. Prevost. They could be feeding on Lomatium,
which occurs plentifully near the summit. In my former paper
on hilltopping F. zelicaon, I gave as my opinion that the F.
zelicaon population on Mt. Arrowsmith were feeding on Loma-
tium. But I later came across a hollow near the summit which
supported a good stand of Heracleiim lanatum, a favorite host
of F. zelicaon. Still, Lomatmm remains a likely host, and the
availability of food plants the most likely theory to account for
the variable numbers of butterflies on different hills.
Shields made no attempt whatever to learn whether his virgin
females could reach the summit if posed any problems in find-
ing it. He did not quite release them on the summit but he
might as well have. Certainly the non-recovery of the mated
females is surprising and must prove something. But it does
not prove that the virgin females went to the summit in order
to find mates, although that would be a reasonable assumption,
if there were not so much evidence against it. My guess is
that if these reared females had been released out of sight of
any hilltop, the virgins would have been recovered close at
hand. The mated females, of course, have a strong urge to
search for a suitable host plant, and this would account for
their moving quickly away from the scene of their release.
If butterflies commonly attempted to reach hilltops from any
distance, one would expect while collecting to note among all
butterflies a cross country movement in a particular direction.
Instead, nearly all of them tend to fly low, and, if they do not
stay in the same place, they travel in such directions as will
not force them to fly over or through trees. This is very
noticeable when collecting on roads, when it is very easy to
intercept one’s quarry, or follow it for long distances, because
of its reluctance to leave the nice clear track.
In the swamp here, female F. zelicaon are usually seen travel-
ling slowly, with a rather hovering flight, just above the
herbage, frequently dropping out of sight therein. Males patrol,
also just above the herbage, evidently on the lookout for females.
Since this quest often brings them down into the sedges, both
8(3):105-117, 1969(1970)
HILLTOPPING
113
sexes exhibit a characteristic damage to their wings, consisting
of numerous small cuts and nicks inflicted by the saw edges of
the sedges. The reader will have remarked that I collected the
summer flight for only about a week. The reason is that these
sedge inflicted abrasions become so prevalent after a short time
that the butterflies are not worth collecting any longer. On the
hill, this type of damage did not show up at all, and I was
able to take saleable specimens for over a month. This again
provides evidence that the small numbers of P. zelicaon on the
hill, in contrast to those in the swamp, are almost certainly
due to the fact that no butterflies from the swamp ever get so
far. Somewhere close to the hill there must be small patches
of a suitable host plant, not associated with sedges.
There is a vast difference between my experience with
Rhopolocera in general, and those of Shields and others, who
list a large proportion of available species as hilltoppers. Part
of this discrepancy, as I have already suggested, may be due to
differences in the general aspect of the terrain. But I still find
it very difficult to accept the idea of possible hilltopping, under
any circumstances, of many species. Among the Lycaenidae, for
instance, there are many species that I never see more than 50
yards away from a good stand of the appropriate host plant.
Mt. Benson offers a particularly good opportunity for asses-
sing the hilltopping proclivities of butterflies. I have visited
this summit an estimated 60 times during the past 24 years.
On each visit I walk about four miles from an elevation of about
2000^ to the summit at 3300^ On this hike, I have collected 30
species of butterflies, of which one, P. zelicaon, is almost always
taken at the summit only, and two others, Vanessa cardui and
V. atalanta, tend to be at the summit more often than elsewhere.
The other 27 species are definitely not more numerous at the
summit, and in many instances are less so. I have not included
Papilio rutulus and P. eurymedon in my count of species, al-
though I have taken both species infrequently in the first part
of the climb. To have included them would have given the
impression that I do not consider them to be hilltoppers, which
they most definitely are. Their absence from Mt. Benson sum-
mit seems to be due to the fact that they have a strictly limited
altitudinal range. It is interesting to note that this aversion to
going beyond a certain height (about 2500' on Vancouver
Island) completely inhibits their hilltopping instinct.
One of the commonest butterflies on Mt. Benson is Oeneis
nevadensis. Females are not commonly seen at the summit.
114
RICHARD GUPPY
J. Res. Lepid.
Males are as plentiful on every little hump of rock or subsidiary
peak, as at the summit. This circumstance does not support
either the theory of Shields (and others) to the effect that the
butterflies head for a conspicuous object on the horizon, or my
theory of insects following ground contours. It seems much
more probable, with O. nemdensis at any rate, that the butter-
flies have never been so far away from these sites selected as
territories that they cannot easily blunder on them by chance.
On my hill at Wellington, the illusion of hilltopping was im-
parted because there were no acceptable rock humps except at
the top. Lately I have come across O. nevadensis males using
as territories patches of bare sandstone showing no eminence
above the plain. Evidently the exposed rock has a considerable
influence on their choice. In the only occasion on which I
have been able to observe an unconfined female O. nevadensis
ovipositing, the act took place right on the summit of the
Wellington hill, again supporting my theory that the insects do
not go far to find their territories.
Several males may occupy the same territory. The very
sparse population of O. nevadensis on my Wellington hill made
it easy to suppose that only one male could remain on a site.
Actually, when an insect has kept a territory to itself for a
short time, any other male arriving will be accosted and perhaps
driven off. But the principle, now well known to zoologists,
that a stimulus applied too often over a short time, will produce
a progressively weakening reaction, applies in this case. When
several males are continually invading a territory, they become
accustomed to one another. They then accost each other only
briefly, and do not fight. This rule applies to other territory
holding butterflies, including the Papilio species.
Limenitis lorquini provides another good example of a terri-
tory-holding species. But the reasons governing this butterfly’s
choice of sites are not nearly so evident as is the case with
others that I have dealt with. After observing a number I have
noticed a similarity. Most consist of a bare or grassy patch on
a south facing slope, with dense shrubbery or trees at the
upper end. The butterflies settle frequently on these shrubs or
trees at varying heights from the ground. It is evident that a
warm air current will travel up the slope to be intercepted by
the trees at the top. I must make it clear that I am not claiming
that most specimens of L. lorquini are found in these situations.
Large numbers are found in what may be makeshift territories,
or may not be territories at all, or the butterflies may be visiting
8(3):105-117, 1969(1970)
HILLTOPPING
115
water, or wet spots to obtain moisture. But when a certain spot
is consistently used by L. lorquini males, when it is always re-
occupied shortly after being cleared off by the collector, then
such a spot will usually fit the above description.
In recent years, the territory holding-habit has come in for
much attention, and it has been shown that it exists in some
degree in a very large proportion of animal species. Many ex-
periments with different animals have shown that they possess
an uncanny ability to return to their home territory, even over
a completely unfamiliar course. But it has been shown that this
ability is not inborn, nor is it necessarily its natal area which
the animal knows as home. An awareness of the territory to
be known as home must become imprinted on the animal, and
this process may take a certain amount of time.
Among animals, winged insects must be particularly likely to
be carried against their will by wind; moreover, their eyes are
not fitted for making out fine detail. It is reasonable to suppose
that insects may have some difficulty in remaining on any
selected spot long enough to become familiar with it, so as to
be able to return from a distance should such a necessity arise.
Conspicuous features of the terrain, such as a hilltop, would
help a lot in obviating this difficulty. Add to this the advantages
of the heat holding qualities of rocks, warm updrafts, and ex-
•posure to the sun early in the day, and I think we have a fairly
good theory to account for the selecting as territories of sites
possessing the several features described above. But I still
remain convinced that the main factor influencing the selection
of a territory is its proximity to the spot where the insect com-
menced its adult life.
By accepting the idea that insects make no great effort to
find a hilltop, but merely use one as a territory if they happen
to blunder on to it, or see it, the objections outlined above are
avoided; in contradistinction we need not suppose that hilltop-
ping can become a blind instinct spread through the population
by natural selection. This would account for some insects using
hilltops which do not appear to offer many favorable features.
Over unusually favorable terrain, such as that so well depicted
in Shields’ illustration, insects might go to a hill by sight from
quite a distance. Shields mentions particularly a marked butter-
fly which reached a hilltop concealed from the release point by
a ridge. But to accomplish this feat the insect required nine
days. Surely, in wandering at random for that length of time,
116
RICHARD GUPPY
/. Res. Lepid.
it is not surprising that it got into a position from which it
could see the hill.
The study of congregations of insects, with a view to proving
that a mating rendezvous is involved, could easily be approached
with too single-minded an attitude. When the primary reason
for the congregation is obvious, as when certain species of
beetles appear in great numbers on a fallen tree, we do not
express any surprise on noting the large number of pairs in
copulation. We know that, in order to reproduce, these beetles
must find a tree of their correct host species in a condition that
makes it vulnerable to their attack. We do not dwell on the
fact that the male beetles can have no interest in the tree
itself, or we may suppose that it is emanations from the female
beetles, rather than the tree, which attracts the males. The last
may be the true explanation but that does not alter the fact that
a knowledge of the beetles’ life history is necessary in order to
evaluate the true reason for the congregation.
To sum up, I consider that hilltopping is usually an aspect
of territorial behavior. With many insects, hilltops provide a
preferred site for territories, and will be used for that purpose
when they can easily be reached from the point where the
insect commences its adult life. When a number of individuals
of a single species reach the same hilltop, they can manage by
splitting it up into small territories, or by sharing a territory.
Explanations to account for territorial behavior can be a very
involved subject. Shields, citing various authors, mentions:
(1) decreased chances of mass predation by a few predators,
(2) less time spent in intraspecific aggression, (3) increased
frequency of male-female encounters, and (4) decreased inter-
ference to courting and mating pairs by other males. Therefore,
hilltopping can be said to facilitate mating to whatever extent
territorial behavior in general facilitates mating.
The above discussion deals with a particular aspect of hill-
topping. Obviously, there are other reasons why insects, in
congregations or singly, are to be found on hilltops. Apart
from species that require arctic or subarctic conditions, which
may be found on mountain tops, there are some that prefer a
hilltop habitat for less obvious reasons. Often they are found
on hills which are not high enough to provide alpine conditions,
but since they are not found in the surrounding area, they
cannot be called hilltoppers. On Vancouver Id,, three species
of Arctiid moths provide good examples of such behavior. They
8(3): 105-1 17, 1969(1970)
HILLTOPPING
117
are Alypia ridingsi Couper, A. langtoni Grt. and Leptarctica
californiae Wlk.
Lastly there is the strange fact that hilltops are always likely
places to turn up unusual locality records. I will not venture
any theory to explain this phenomenon. From a number of my
own interesting hilltop captures, both in the Lepidoptera and
the Coleoptera, I will select the most remarkable as an example.
I refer to the taking, on Mt. Arrowsmith in August of 1966, of
a specimen of Fieris sisymbrii Bdv. The specimen was sent to
Dr. dos Passos for positive identification. The species was not
previously known to exist anywhere west of the coastal moun-
tains in British Columbia. This individual had enough of the
wing area torn off, on one side only, to seriously impede its
flying ability.
LIST OF RHOPALOCERA SPECIES COLLECTED ON
MT. BENSON
Subspecific names omitted
Papilio zelicaon
Farnassius clodius
C olios occidentalis
Neophasia menapia
Cercyonis alope
Oeneis nevadensis
Speyeria hydaspe
Boloria epithore
Polygonia faunus
F zephyrus
Nymphalis milberti
Vanessa atalanta
V. cardui
Limenitis lorquini
Strymon melinus
S calif ornicus
Incisalia iroides
L fotis
I eryphon
Lycaena mariposa
L helloides
Everes amyntula
Plebeius melissa
P icarioides
Glaucopsyche lygdamus
Thorybes pylades
Pyrgus ruralis
Erynnis icelus
Hesperia harpalus
Ochlodes sylvanoides
Journal of Research on the Lepidoptera
8(3):118-128, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
IDENTITY OF THE MOTH LOXAGROTIS
PAMPOLYCALA (DYAR) FROM THE
SOUTHWESTERN UNITED STATES AND MEXICO
(NOCTUIDAE)
JOHN S. BUCKETT
Department of Entomology
University of California, Davis
Over the past two decades or more, a species of Loxagrotis
McDiinnough near L, socorro (Barnes) has been collected in
southern Arizona. It was thought to represent a new species, and
only recently the author checked the types of similar species,
including the type of L. pampolijcala (Dyar), in the United
States National Museum. The type male of pampolycala,
U.S.N.M. type number 14210, described from Mexico, matches
the series of the males before the author in every detail.
At the time of the original description of pampolycala, Dyar
(1912) had only two specimens before him, a male and a female.
In his discussion immediately following his original description,
Dyar stated concerning the two specimens before him “The
female before me is similar [to the male] . . The brackets are
mine. From this statement, it seems obvious that the male should
be selected as lectotype at the time of revisionary work.
McDunnough (1928) in his “A Generic Revision of North
American Agrotid Moths” stated (p. 28) concerning four species
of Loxagrotis “For the present salina Barnes, capota Sm.,
albicosta Sm., and socorro Barnes are placed here although the
latter two are quite atypical, the palpi being heavily but smooth-
ly scaled, not fringed with hair, and the genitalia of each show-
ing a complete corona and considerable individual difference in
the position and shape of the harpe.” It is well to note that
pampolycala also should be placed with this “atypical” group of
the genus, namely with albicosta (Smith), capota (Smith), salina
(Barnes) and socorro (Barnes).
118
8(3) ;1 18-1 28, 1969(1970)
LOXAGROTIS
119
Fig. 1. Loxagrotis pampolycala (Dyar), male. Madera Canyon, Santa Cruz
County, Arizona, 7 July 1963, Bauer-Buckett slide no. 69B25-33
( W. R. Bauer & J. S. Bucket! ).
Fig. 2. L. pampolycala, female. Data same as in fig. 1.
120
JOHN S. BUCKETT
/. Res. Lepid.
The synonymy of pampolijcala under socorro is due to the
work of Draudt (in Seitz, 1923). He states (op. cit. ) “f. pampoly-
cala Dyar belongs hereto, it is marked exactly the same, but
more iron-grey in the ground-colour, without the dark filling of
the cell and not so variable. The hindwing of the female is like
that of the male whitish and hardly darkened. Mexico.” The
hindwing of the female, however, is somewhat darkened and can
hardly be called “white”.
Barnes and Benjamin (1924) followed this synonymy of pam~
polijcala beneath socorro, however considered both species to
belong to Chorizagrotis Smith, rather than to Rhizagrotis Smith
as did Draudt. McDunnough (1928), was then the first author
to place the concerned species in his newly erected genus
Loxogrotis.
Loxa grot is pampolijcala (Dyar)
Lijcophotia pampolijcala Dyar, 1912, Proc. U.S. Nat. Mus.
42:57-58.
Rhizagrotis socorro, form pampolycala, Draudt {in A. Seitz),
1923, Macrolepidoptera of the World, 7:51.
Chorizagrotis pampolycala, Barnes and Benjamin, 1924 (as a
synonym of socorro), Contrib. Nat. Hist. Lepid. N. America
5(3):111.
Loxagrotis socorro, form pampolycala, McDunnough, 1938,
Mem. Southern California Acad. Sci. 1:61.
DESCRIPTION: Male: Head with vertex and Irons evenly
rounded, slightly roughened, protruding; palpi exterolaterally
blackish, basal segment colored ventrally with tan colored elon-
gate hairs; apical portion of second segment light tan, third
segment stubby, colored in tan scales; compound eyes with band
of blackish hairs exterolaterally; antennae with scape and pedicel
clothed in flattened tan colored scales; flagellomeres dorsally
clothed with tan colored scales, ventrally fasciculate, apically
becoming setose-ciliate. Thorax with collar composed of elongate
dentate scales, basally tan, subapically somewhat darker, tan
tipped; tegulae composed of flattened elongate tan colored
scales and elongate brown simple hairs; disc composed pre-
dominently of elongate tri-colored flattened hairs, basally tan,
subapically brown, apically tan; ventrally clothed in elongate
tan colored hairs; tarsi with segments clothed in black scales
8(3);118-128, 1969(1970)
LOXAGROTIS
121
Fig. 3. L. socorro (Barnes), male. Sunnyside, west side, Huachuca Moun-
tains, Cochise County, Arizona, 9 July 1958 (L. M. Martin).
Fig. 4. L. socorro, female. Madera Canyon, Santa Rita Mountains, Santa
Cruz County, Arizona, 10 July 1957 (L. Stange and Harding).
122
JOHN S. BUCKETT
J. Res. Lepid.
except for apical annuli of tan colored scales; primaries with
basal line represented costally as dark brown mark, thence by
a single black dot on second anal vein; transverse anterior area
with a longitudinal black elongate medial band; transverse
anterior line hardly discernable and when visible appearing
geminate, basally tan apically dark brown, course as in figure 1;
medial area tan, overlain with dark brown scales; orbicular longi-
tudinally elongated, tan, blending into tan costal band; reniform
ovate, composed of light tan scales, outlined in dark brown
scales, these scales coalescing with dark brown outline of orbic-
ular; transverse anterior line scalloped, dark brown to black,
course as in figure 1; subterminal area costally dark brown,
thence tan colored, overlain with dark brown scales to inner
margin, veins ventrally outlined in darker color; subterminal line
very irregular, represented basally in dark brown, thence a band
of tan scales terminally; terminal line composed of dark brown
scallops between veins, these scallops being very shallow; fringes
tri-colored, basally tan, medially dark brown, remainder off-
white; ventral surface tan, with a suggestion of transverse pos-
terior line in dark brown; secondaries whitish with a bluish tinge,
costally tan colored; terminal line dark brown, fringes white;
ventral surface as in dorsal surface except for presence of dark
brown exterior band on costa. Abdomen dorsally clothed in
elongate tan colored scales and simple hairs; ventrally clothed
in tan colored scales and hairs which overlay broadened simple
white colored scales. Greatest expanse of forwings 17 mm.
Genitalia as in figures 5 and 6.
Female: As in male except antennae ciliate; secondaries dirty
whitish overlain with brownish scales appearing almost fuscous,
as in figure 2. Greatest expanse of forwings 18 mm. Abdomen
dorsally clothed in brown broadened scales and simple hairs,
posterior portion of segments clothed in light brown; ventrally
clothed in off-white scales and hairs. Genitalia as in figure 9.
SPECIMENS EXAMINED: Mexico: Cotypes, no. 14210, U.S.
Nat. Mus., 1 male, Cuernavaca, May, 1911 (R. Muller); 1 female,
Guerrero, Mexico (J. Doll). Arizona: 1 male, 3 females, Madera
Canyon, Santa Cruz County, 4880' elevation, 7 July 1963 (W. R.
Bauer & J. S. Buckett); 2 females, same data as preceding, 8
July 1963; 1 female, Madera Canyon, Santa Cruz County, 16
July 1967 (C. W. Baker).
Specimens studied are deposited in the Entomology Depart-
ment, University of California, Davis and the collection of the
8(3);118~128, 1969(1970)
LOXAGROTIS
123
Fig. 5. L. pampolycala, male. Genitalia minus aedeagus. Data same as in
Fig. 1.
Fig 6. L. pampolycala, male. Aedeagus, inflated, data same as in fig. 5.
124
JOHN S. BUCKETT
/. Res. Lepid.
Fig. 7. L, socorro, male. Genitalia minus aedeagus, Madera Canyon, Santa
Cruz County, Arizona, 6 July 1963, Bauer-Buckett slide no.
69B25-31 (W.R.B. & J.S.B.) .
Fig, 8. L. socorro, male. Aedeagus, inflated, data same as in fig. 7.
8(3):118~128, 1969(1970)
LOXAGROTIS
125
Fig 9. L. pampolycala, female genitalia. Madera Canyon, Santa Cmz
County, Arizona, 8 July 1963, Bauer-Buckett slide no. 69B25-34
(W.R.B. & J.S.B.).
126
JOHN S. BUCKETT
J. Res. Lepid.
Bureau of Entomology California Department of Agriculture,
Sacramento.
Loxagrotis pampolycala differs from its closest relative, L.
socorro by being slightly larger, and more drab in coloration.
Also, the reniform of pampolycala is larger and more in a
diagonal position on the primaries, whereas the reniform of
socorro is more upright on the wing. The secondaries of the
females of pampolycala are darker than are those of socorro too.
Both species occur sympatrically, and to my knowledge, nothing
is yet known concerning the life histories of either species.
Loxagrotis socorro ( Barnes )
Rhizagrotis socorro Barnes, 1904, Canad. Entomol. 36(6) :171-
172; Barnes and McDunnough, 1912, Contrib. Nat. Hist. Lepid.
N. America 1(4): 16, pi. 6, fig. 20; Draudt, (in A. Seitz), 1923,
Macrolepidoptera of the World 7:51.
Chorizagrotis socorro, Barnes and McDunnough, 1917, Check-
list of the Lepidoptera of Boreal America, p. 44; Barnes and
Benjamin, 1924, Contrib. Nat. Hist. Lepidoptera of N. America
5(3):111-112.
Loxagrotis socorro, McDunnough, 1928, Canad. Dept. Mines,
Bull. no. 55, Biological series no. 16:27-28; 1938, Mem. Southern
California Acad. Sci. 1:61.
DIAGNOSIS: Vestiture of head brown to dark brown; an-
tennae of male fasciculate, terminally becoming setose-ciliate, of
female ciliate. Thorax with divided collar possessing a dark
transverse band; disc and tegulae clothed in various shades of
brown; ventrally clothed in elongate whitish simple hairs; pri-
maries dorsally with maculation as in figs. 3 and 4, ground color
dark brown; costa with conspicuously cream colored band, from
base to just past reniform; reniform ochreous, centrally filled with
dark brown scales; subterminal area conspicuously washed with
whitish scales, contrasting with median and dark brown terminal
areas; secondaries whitish with purplish sheen in male, in female
there is tendency toward dirty white or fuscous. Greatest expanse
of forewings 15-16 mm. Genitalia as in figs. 7, 8, and 10.
SPECIMENS EXAMINED: Arizona: 1 male, Madera Canyon,
Santa Cruz County, 4880' elevation, 6 July 1963 (W. R. Bauer
& J. S. Buckett); 2 females, same data as preceding, 7 July 1963;
1 female, same data as preceding, 8 July 1963; 1 female, same
data as preceding, 14 July 1963; 1 female, Madera Canyon, Santa
Fig. 10. L. socorro, female genitalia. Madera Canyon, Santa Cruz County,
Arizona, 7 July 1963, Bauer-Buckett slide no. 69B25-32 (W.R.B.
& J.S.B,).
128
JOHN S. BUCKETT
/. Res. Lepid.
Rita Mountains, Santa Cruz County, southern Arizona, 10 July
1957 (Stange and Harding); 1 male, Sunnyside, west side
Huachuca Mountains, Cochise County, ex. 15 watt fluorescent
black light, 8 July 1958 (Lloyd M. Martin); 1 female, same data
as preceding, 9 July 1958; 1 male, same data as preceding, 12
July 1958.
L. socorro may be distinguished from its closest relative, L.
pampolycala by use of the characters in the discussion section
under pampolycala, as well as by use of the genitalia.
LITERATURE CITED
BARNES, WM., 1904. New species of North American Lepidoptera. Canad.
Entomol 36(6) :165-173.
, and J. H. McDunnough, 1912. Contributions to the natural
history of the Lepidoptera of North America, Illustrations of rare and
typical Lepidoptera I (4): 1-62, inch 27 pis., the Review Press,
Decatur, Illinois.
, and J. H. McDunnough, 1917. Checklist of tlie Lepidoptera
of Boreal America. Herald Press, Decatur, Illinois, 392 -j- vii pp.
, and F. H. Benjamin, 1924. Contributions to the natural
history of the Lepidoptera of North America, notes and new species.
5(3):1-199, the Review Press, Decatur, Illinois.
DRAUDT, M. (in A. A. SEITZ), 1923. The Macrolepidoptera of the
World. Alfred Kernen Press, Stuttgart, vol. 7, 412 pp. -j- 96 pis.
DYAR, H. G., 1912. Descriptions of new species and genera of Lepidoptera,
chiefly from Mexico. Froc. U.S. Nat. Mus. 42:39-106.
McDunnough, j. H., 1928. a generic revision of North American Agrotid
motlis. Canad. Dept. Mines, Bull. no. 55, biological series no. 16, 78 pp.
, 1938. Check list of the Lepidoptera of Canada and the
United States of America, part 1, Macrolepidoptera. Mem. Southern
California Acad. ScL, vol. 1, 272 pp.
Journal of Research on the Lepidoptera
8(3):129-132, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
RECORDS OF
COLIAS GIGANTEA STRECKER (PIERIDAE)
FROM SOUTHEAST MANITOBA
AND ? MINNESOTA
JOHN H. MASTERS '
P. O. Box 7511, St. Paul, Minnesota
HoVANITZ (1950) RECORDED THE KNOWN RANGE of ColiaS
gigantea Strecker as subarctic North America, from the northern
limit of trees in Alaska east to Hudson Bay; southward, in willow
bogs, to southern Canada, from the Rockies in Alberta eastward
to the Manitoba Escarpment; then on south in the Rockies (as
scudderii Reakirt) to Colorado and New Mexico — but suggests
a possible range extension eastward to northern Minnesota,
Wisconsin and Michigan in willow bogs. Up until now the only
recorded range extension recorded was by Riotte (1962) who
recorded specimens from three points on Hudson Bay in Ontario
that extended the range eastward from Churchill to Fort Albany
on James Bay. John Polusny, of Winnipeg (in litt. ) reports a few
solitary specimens of Colias gigantea in southeast Manitoba;
but it is only recently that I have been able to confirm that it
occurs south of the 53rd parallel, east of Lake Winnipeg.
On 8-10 August, 1969, I collected 5 males and 2 females of
Colias gigantea at three localities between the O’Hanley and
Sand (sometimes called Sandy) Rivers along Manitoba highway
304. These locations are just east of Lake Winnepeg and are
between the 50th and 51st parallel. All of the specimens were
taken in wet sphagnum/ willow bogs and all were somewhat
worn in condition. The determination of the females leaves
little room for doubt: one of them is “white” with wing borders
completely immaculate of black (this is a common variant situ-
^ Research Associate, Section of Insects and Spiders, Carnegie Museum,
Pittsburgh, Pennsylvania.
129
130
JOHN H. MASTERS
/. Res. Lepid.
Fig. 1. Probable CoUas gigantea Strecker. Male taken 2 August 1967 by
W. A. Bergman near McNair, Lake County, Minnesota. Upper side above,
lower side below. 175 percent actual size.
8(3):129~132, 1969(1970) COLIAS GIGANTEA
131
ation with female gigantea) while the other female is yellow
with fully developed, but weakly scaled, black borders. The
five males are distinguished from C. interior (with whom they
are superficially very similar) by their larger size and association
with willow bogs and the indicated females. A few specimens
of Colias interior Scudder were taken in nearby locations. They
were quite worn and tattered, indicating that C. interior prob-
ably has an earlier flight here than C. gigantea. The south-
eastern Manitoba specimens of Colias gigantea are very close in
appearance to Colias gigantea mayi Chermock & Chermock from
Riding Mountain, Manitoba. Like mayi they lack the heavy
dark scaling on the ventral hindwing that is usually used as an
identification character for Colias gigantea. Because of this, the
key to Colias by Klots ( 1961 ) is ineffective for separating them
into gigantea — the confusion resulting at couplet 4a-4b.
On August 2nd, 1967, Bill Bergman of Minneapolis collected
a very large male Colias at McNair, Lake County, Minnesota.
He was collecting along a railroad track that borders a bog
when he noticed the butterffy in flight because of its apparently
large size. After mounting the butterfly he asked me to examine
it thinking it might be C. gigantea. In Colias a definitive de-
termnation of a single specimen is often impossible and until
more Minnesota examples are collected we cannot be 100%
positive that this specimen is indeed gigantea. However, in my
own mind I am fairly certain that it is gigantea. The butterfly
was taken in a potentially suitable habitat for gigantea, has a
large wing expanse (29 mm. base to tip of forewing which is
larger than any specimen of C. interior that we have seen from
Minnesota), and while the specimen is quite worn, the mark-
ings (see figure 1) seem to fit in better with a series of gigantea
than they do with interior. There are a couple of large male
Colias in the John Nordin Collection, at Webster, South Dakota,
which were taken in Koochiching County, Minnesota that also
might prove to be gigantea. The Bergman specimen from Lake
County has been placed in the American Museum of Natural
History in New York City.
In addition to the specimens of C. gigantea mentioned in the
preceeding, I have taken several dozen specimens along the
Manitoba Escarpment from Riding Mountain northward in
western Manitoba. My preliminary field data suggests that the
species may be associated with string bogs. String bogs are
peculiar in that they are topographically aligned in strips up
132
JOHN H. MASTERS
/. Res. Lepid.
and down slope. The climatic and edaphic conditions necessary
to produce string bogs is not understood, but it is clearly differ-
ent from the lake fill succession that is normally accepted for
bogs south of the 53rd parallel. Most bogs north of the 53rd
parallel are a result of the underlying permafrost that prevents
water drainage through the top soil. String bogs, by nature of
their being topographically aligned, can be assumed to have
groundwater movement which may favor, at least in some
areas, the growth of willows over black spruce. Heinselman
(1963) records the presence of string bogs as far south as the
north shore of Red Lake in Minnesota and ( 1965 ) at an isolated
spot near Seney, Michigan. It would be interesting if the range
of C olios gigantea continues to correspond with the areas in
which string bogs occur as both of them become better known.
LITERATURE CITED
HEINSELMAN, M. L., 1963. Forest sites, bog processes, and peatland
types in the Glacial Lake Agassiz region, Minnesota. Ecol. Mono-
graphs 33: 327-347.
1965. String bogs and other patterned organic terrain near
Seney, Upper Michigan. Ecology 45: 185-188.
HOVANITZ, W., 1950. The biology of Colias butterflies. I. The distri-
bution of the North American species. Wasman Jour. Biol. 8: 49-75.
KLOTS, A. B., 1961. Genus Colias in Ehrlich, P. R. & A. H. Ehrlich.
How to know the butterflies. Wm. G. Brown Go., Dubuque, Iowa,
262 pp.
RIOTTE, J. G. E., 1962. First additions to the northern Ontario list of
butterflies. Jour. Lepid. Soc. 16: 243-245.
BOOKS:
NOTICES
BUTTERFLIES. A concise guide in colour. Josef Moucha, ill. by
Vlastiinil Choc. Paul Hamlyn, Hainlyn House, The Centre,
Felthain, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper back reprint, N.Y,
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOCiY. Theodore H. Savory. Philosophical
Library, N.Y.
WANTED:
Brephidium exilis, B. fea, B. isophtfialma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Ave., Arcadia, Galifornia 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systeniatics, biogeographical and genetic approach to an under-
standing of this group of inseets.
NEEDED:
Manu.scripts for immediate publication in (his JOURNAL. With color
nlay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3 \ 5 card in the form desired and they will be printed in the
next following issue of the JOURNAL.
Volume 8
Number 3
September, 1969
IN THIS ISSUE
Development of the markings on
the pupal wing of Pieris pupae.
John M. Kolyer 69
Two new forms of Plebejinae from Wyoming. 91
Concerning the names and status of
certain North American members of the
genus Phycoides. J. W. Tilden 94
The life history of Amblyscirtes linda
( Hesperiidae ) .
J. R. Heitzman and Roger Heitzman 99
Further observations on “hilltopping” in
Papilio zelicaon. Richard Guppy 105
Identity of the moth Loxagrotis
pampolycala ( N octuidae ) .
John S. Buckett 118
Records of C olios gigantea from
southeast Manitoba and Minnesota.
John H. Masters
129
Volume 8
Number 4
December, 1969
THE JOURNAL OF RESEARCH
©NJ THE LERIJDOFTERA
published by
The Lepidnptera Research Foundation, Inc.
at
1 160 W. Oranj^e Grove Avc., Arcadia, Calif. U.S.A. 91006
EDITOR: William Hovanitz
Associate Editors :
I'homas C. Emmel, Dept, of Zoology, University of Florida, Gainesville,
Florida 32601.
Maria Etcheverry, Centro de Estiidios Entomoloj^icos, Casilla 147, Santiago,
Chile.
T. N. Freeman, Div. of Entomology, Dept, of Agriculture, Ottawa, Ontario,
Canada.
Brian O. C. Gardner, 18 Chesterton Hall Crescent, Cambridge, England.
Rudolf H. T. Mattoni, 9620 Heather Road, Beverly Hills, Calif. 90210.
Lee D. Miller, The Allyn Foundation, Inc. Sarasota Bank Building,
Sarasota, Florida 33578.
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Journal of Research on the Lepidoptera
8(4):133=152, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
SEASONAL CHANGES IN ORGANIZATION OF
TROPICAL RAIN FOREST
BUTTERFLY POPULATIONS IN PANAMA
THOMAS C. EMMEL and CHARLES F. LECK
Department of Zoology, University of Florida, Gainesville 32601
Section of Neurobiology and Behavior, Cornell University, Ithaca 14850
Seasonality is usually considered a distinctive feature of
temperate zone communities, with their fluctuating annual cli-
mates, while the wet lowland tropics are relatively constant in
most environmental conditions. Yet major seasonal changes in
tropical plant communities have been demonstrated for even
rain forest areas and the selective reasons for this seasonality
are beginning to be explored (e.g., see Janzen 1967). The
possible existance of seasonality in the animals of tropical com-
munities has barely been touched upon to date, most available
data being on birds (e.g., Skutch 1950, Moreau 1950, Miller 1954,
Ricklefs 1966, Leek 1970), certain tropical lizards (Hirth 1963,
Alcala 1966, Sexton 1967) and foliage-inhabiting insects (Janzen,
unpublished; Hespenheide, unpublished). These studies have
shown there may be significant changes in vertebrate population
size and reproductive activities even in tropical forests with a
constant annual eflmate. Differences in population density and
species composition of foliage-inhabiting insect communities have
been shown (by Janzen and Schoener 1968) between wetter and
drier sites in a tropical deciduous forest, while the dry season
has been shown to decrease population density of inflorescence-
feeding Drosophila in Panama (Pipkin, Rodriguez and Leon
1966). Most mosquito species studied by Bates (1945) in tropi-
cal eastern Colombia showed seasonal fluctuations in population
density. Yet there has been little documentation of the recent
textbook assertion (Boughley 1968: p. 40) that “Marked seasonal
fluctuations in population density are encountered as frequently
134
EMMEL AND LECK
J. Res. Lepid.
Fig. 1. — Map of Barro Colorado Island, Canal Zone, showing location of
1968-69 study areas (Laboratory Clearing and Forest trail).
8(4):133-152, 1969(1970) SEASONAL CHANGES
135
in tropical . . , regions as they are in temperate.” Moreover,
detailed data on seasonal changes in species composition of a
major animal group aside from birds (Slud 1960) are lacking
for the tropical rain forest in the Americas.
The existance of seasonal change in species composition, as
well as number of individuals, is a standard feature of temperate
communities, especially among the arthropods but even ver-
tebrates (e.g., birds). One group of insect species is character-
istic of the spring fauna, another group of species replaces them
in early summer, and so on with the changing temperature
regime and food availability. However, the tropical species in
rain forests would be expected to breed all year if constant tem-
perature was the principal requirement for continuous breeding.
A stable species composition in any one area should result; that
is, the same group of species should be present all year. On the
other hand, fluctuations in population density and even species
composition could be expected from possible seasonal variation
of rainfall, humidity, light intensity as affected by cloudiness,
and other environmental conditions in the rain forest.
With about 4,000 species in tropical America (Seitz 1913) out
of a world fauna of 12-15,000 species and their suggested impact
upon evolution of the angiosperms (Ehrlich and Raven 1965),
butterflies definitely qualify as a major tropical animal group
of considerble ecological interest. This study examines the dy-
namics of faunal composition in the resident butterflies of the
tropical rain forest in Barro Colorado Island, Panama, with
respect to seasonal changes in active species and changes in
population density from the latter half of the wet season
(October-November 1968) through the major portion of the dry
season (December 1968-March 1969). The viewpoint that
tropical species diversity may be influenced by a “seasonal eco-
tone” or edge effect at the period of wet-to-dry-season transition
is suggested by these faunal changes in Panamanian butterflies.
DESCRIPTION OF STUDY AREAS AND METHODS
This research was conducted on Barro Colorado Island in
Gatun Lake, Canal Zone, Isthmus of Panama, from October 1,
1968, through April 12, 1969. The island is largely covered
with rain forest having a canopy starting at about ninety feet,
The annual rainfall is about 2700 mm at the Smithsonian Trop-
ical Research Institute station (Moynihan, 1968). Further de-
scriptions of the general vegetation and climate may be found in
AUee (1926a, 1926b).
136
EMMEL ANE LECK
/. Res. Lepid.
SPECIES
PAPILIONIDAE
Papilio anchisiades
P- arcus
P. •rftholon
P. gloucolaus
P. polydamus
P. thoas
PIERIDAE
Appias drutillo
Cotopsilio statira
Euremo daira
E. lisa
E. marginella
E. messalina
Itaballia demophile
I. pisonis
P«rrhybris pyrrha
WET SEASON DRY SEASON
Fig. 2.— Patterns of flight activity of adult populations of butterflies in
the Families Papilionidae and Pieridae, Clearing study area, October 1968
to the end of February 1969. The width of the bar indicates relative adult
population density for a given species during each month (number of
days species was observed out of total number of observation days).
8(4):133-152, 1969(1970) SEASONAL CHANGES
137
Clearing Study Area. Most of the data reported herein were
collected in the clearing extending from slightly southwest of
the Smithsonian Tropical Research Institute field station north-
eastward to the shore of Gatun Lake, at 9°09'50" north latitude
and 79°50'25" west longitude (see Fig. 1). This clearing mea-
sures about 778 m by approximately 286m (the width varies
considerably ) ; the elevation at the southwestern end is 35 m
higher than the northestern end at the Gatun Lake shore line
(26m above sea level). Since its establishment, the size and
condition of the clearing has varied considerably through the
years. Presently it is maintained in the early stages of succession,
much as described by Kenoyer ( 1929 ) : ( 1 ) In small areas about
the buildings frequent cutting permits only grasses and annual
weeds. (2) In much of the clearing where cutting occurs in-
frequently, plants of the second-year association are common
(e.g. Heliconia and Piper). (3) Along the edges and in neglect-
ed patches throughout the clearing, species of the ‘pioneer
forest” (e.g. Cecropia, O chroma, and Tetracera) dominate.
(4) Within the whole area introduced plants are important
(e.g. Citrus, Hibiscus, Ixora, Musa, Psidium, and others).
Gensusing of butterfly species was carried out for two hours
daily on census days: one hour between 0900 and 1200, and
one hour between 1300 and 1500. Specific hour periods were
rotated regularly. Only 4 lycaenid and 2 hesperiid species were
censused; otherwise, all species present were recorded. Field
identification of flying butterflies was considered quite accurate
(all species identifications were originally verified by collected
specimens). All censusing was done by the second-named
author (G.F.L. ).
Forest Study Area. Within the rain forest species censuses
were conducted weekly to record changes in forest-restricted
butterfly populations. The 2,050 m trail route indicated in Fig-
ure 1 was censused from 0800 to 1030; no afternoon censuses
were taken. This route ran west of the C.G.I. Laboratory on
the Lathrop trail, then south and east on the Miller, Wheeler,
and Snyder-Molino trails, and north on the last-named trail to
the laboratory again.
RESULTS
The Clearing study area provided census data on 92 species
of butterflies; these are summarized in Table 1. The relative
abundance or relative population density of each species per
138
EMMEL AND LECK
J. Res. Lepid.
TABLE 1
Flight activity patters in wet and diy seasons for butterfly species in
Clearing study area, October I968 through February 1969. Figures represent
percentage of total census days in a month that each species was recorded.
Relative Adult Population Density of Species*
FAMILY Species WET SEASON DRY SEASON
October November December January February
PAPILIONIDAE: 6 species
Papilio anchisiades
u
h
P. arcus
58
22
20
--
lU
P. erithalon
25
9
5
13
27
P. glaucolaus
—
9
P . polydamus
12
5
-
11
P. thoas
IT
9
lU
35
68
PIERIDAE: 10 species
Appias drusilla
26
Ih
Phoebis statira and argante
U2
Ih
82
87
77
Eurema daira
""
--
9
E. lisa
75
61
77
87
6h
E. marginella
31
55
12
5
E. messalina
9
5
17
—
Itaballia demophile
50
83
86
13
18
!• pisonis
—
--
--
18
Perrhybris pyrrha
57
68
13
23
DANAIDAE: 21 species
Dabaus gilippus
u
....
5
Lycorea cleobaea
8
8(4)^33-152,1969(1970) SEASONAL CHANGES
Table 1: Continued
Oct .
Nov.
Dec .
Jan .
Feb
ITHOMIIDAE: 6 species
Aeria eurimedia
-
—
5
—
11
Hypoleria libera
-
—
5
—
—
Hypothyris euclea
—
h
5
1|
-
Mechanitis franis
—
—
9
—
M. isthmia
—
-
—
35
9
Tithorea tauracina
—
—
—
13
-
SATYRIDAE: 10 species
Antirrhaea miltiades
—
-
5
—
—
Callitaera menander
IT
IT
9
k
—
Euptychia antonoe
—
—
—
9
Ik
E, gulnare
li
—
5
13
5
E. hermes
96
8T
100
8T
k^
E. besione
h
9
5
li
—
E. juani
li
-
—
9
E. labe
—
—
5
—
—
E. molina
T5
U3
53
65
68
Pierella luna
13
IT
Ih
9
18
BRASSOLIDAE: species
Caligo sp. 8 —
Eryphanis polyxena — U
Orsiphanes fabricii
0. xanthicles 13 ^
MORPHIDAE: 3 species
Morpho peleides and amathonte 50 UU
M. theseus —
5
5
5
111
18
13 9
k
39
IT
139
50
140
EMMEL AND LECK J- Lepid.
Table 1: Continued
Oct .
Nov.
Dec .
Jan.
Feb.
HELICONIIDAE: 13 species
Colaenis (Dryas) julia
h2
52
91
70
61t
Dione jxmo
29
13
9
—
—
D. vanillae
—
—
—
—
5
Heliconius (Eueides) aliphera
—
13
—
9
—
H. (E.) isabella
IT
—
-
It
27
H. (E.) lybius
—
—
—
it
—
Heliconius cydno
70
68
50
59
H. doris
-
-
5
30
5
H. erato
88
70
77
78
86
H. ethillius
58
35
32
35
17
H. sappho
-
—
-
9
H . s ara
71
70
68
57
23
Metamorpha dido
h
-
—
-
—
NYMPHALIDAE; 21 species
Adelpha iphicleola
—
U
5
It
5
A. marc i a
8
k
Ih
22
9
Ageronia (Hamadryas) februa
U
13
-
—
5
Anartia fatima
100
100
100
100
100
A. jatrophae
71
13
23
Ik
91
Callicore sp.
-
It
5
—
Catagramma sp. (peralta?)
—
—
—
C. titheas
-
1+
—
h
—
Catonephele numilia
-
9
5
—
-
Myscelia cyaniris
—
9
—
5
Phyciodes clio
5
8(4)^33^152,1969(1970) SEASONAL CHANGES
Table 1: Continued
Oct .
Nov
P. leucodesma
8
P. ofella
--
Precis lavinia
--
k
Prepona sp.
—
Protogonius fabius
13
—
Pyrrhogyra crameri
--
—
Taygetis uncinata
8
—
Temenis libera
U
k8
Marpesia chiron
8
—
Victorina steneles
--
h
RIODINIDAE: 11 species
Caleph'elis virginiensis
--
k
Charis chrysus
k
Eurybia patrona
—
Euselasia sp.
--
—
Hades noctula
—
Ithomeis eulema
—
Mesosemia sp.
U
—
M. telegone
13
13
Nymula phy Ileus
--
Oleria paula
Zelotaea pellex
--
--
LYCAENIDAE: k species tallied
Strymon yojoa
Theda hemon
—
--
T. jalan
__
__
Dec. Jan. Feb.
9 1|
9
U
5 U
9 5
27 30 Ih
9 IT
27 26
5
5
9
k
9
5
k
U
1+
141
142
EMMEL AND LECK
/. Res. Lepid.
Table 1; Continued
Oct. Nov.
T. togarna
HESPERIIDAE: 2 species tallied
Eudamus sp. — U
Hesperia syrichtus 50 13
Dec .
5
32
Jan. Feb.
Ik
65 27
52 77
TOTAL; 92 species tallied in Clearing
*The index of relative population density of each species is recorded
as the frequency of occurrence out of the total number of census days each
month (see text). The numbers of census days per month, I968-69, were:
October (2U), November (23), December (22), January (23), February (22).
The wet season extends from June to mid December, the dry season from late
December to May (see text).
Fig. 3.— Pattern of relative flight activity in the Clearing and Forest areas,
of the pierid butterfly, Itaballia demophile.
8(4):133~152, 1969(1970) SEASONAL CHANGES
143
month is indicted as a percentage:
Number of days sp. recorded Index of Relative Abundance
— — ~ = (Percentage of census days
Total no. of census days that month each sp. was recorded).
That is, the commoner a species the higher the probability that
it will show up in all census periods. For example, Anartia
fatima (Nymphalidae) was the only species observed on all
census days every month (index value of 100%), while Papilio
anchisiades was only seen on one out of 23 days in November
(index =: 4%) and one out of 22 census days in February (Table
1); hence the latter species’ population density was compara-
tively very low. The Clearing area was too large and time too
limited for capture-recapture determinations of absolute popu-
lation densities. However, the present method at least allowed
an accurate estimate of variations in adult population density
from month to month. The data for the period from October
1968 to February 1969 were collected on a comparable number
of days (22 to 24; see Table 1). Only five census days were
available for March 1969, and the data are not tabulated here
though they support the same general trends already evident in
the dry-season censuses.
The Forest study area supported a much smaller fauna;
census data on the 23 species observed are given in Table 2.
Here, actual numbers are given because of the variable number
of census days per month and the low forest population densi-
ties which made sampling errors relatively more important.
The average rainfall and duration of wet and dry seasons for
the last forty years on Barro Colorado Island are given in
Table 3. The rainfall during the present study, September 1968
to March 1969, is given in Table 4. It is clear from a compari-
son of the two tables that the dry season began somewhat earlier
than usual in 1968-69, but that December is a transition period
between the end of the heaviest rains (in November) and the
start of the dry season which come towards the end of December.
The data in Table 5 indicate that our censusing procedure
included a full representation of the butterfly fauna of Barro
Colorado Island, with the exception of the two groups of small,
often secretive or fast-flying species in the families Lycaenidae
and Hesperiidae.
TABLE 2. Flight activity patterns in wet and dry season for butterfly populations
144
EMMEL AND LECK
/. Res. Lepid.
VO on
rH CM rH rH
H
H CM H CM
O
o a
C O
-H
>5 W
O (V
rd
m
C 4::
o o
o >>
•H -p
H ft
C3 O
0 ^
><! -H ft cd
G ft -H
H cd cd T)
o w OJ
ft •'-3 B
0
o
a
• o
ft -p
w a
CO
CO cd jd
0 -H -H
cd o o
0
ft -p -H
•H ft H
Jh Id 0
W H m
t ^
0 -p
< M
•H ?H
H 0
Id 01
•1-3 -H
rH
CO
•H CO
G -H
^Species arranged in order of appearence during forest-survey period, not in taxonomic groups.
8(4):133~152, 1969(1970) SEASONAL CHANGES
145
TABLE 3. Rainfall in wet and dry seasons on Barro Colorado Island, Canal
Zone, Isthmus of Panama (Station average, 1925 or 1926 to 1966;
data calculated
from Moynihan, 1968).
Month
Station Average
Rainfall in mm.
WET SEASON
May
2T6.6
June
2T6.9
July
293.9
August
329.7
September
262.6
October
3UT.5
November
U61.3
December
269.2
DRY SEASON*
January
57. T
February
32.5
March
29.2
April
88.
ANNUAL TOTAL
2,712.7
DRY SEASON; TOTAL:
207.8
WET SEASON: TOTAL:
2,50ii.9
^Dry season generally starts in latter half of December. The median
date for the beginning of the dry season is about December 20.
146
EMMEL AND LECK
/. Res. Lepid.
Table k
Rainfall in the I968-69 study period on Barro Colorado Island,
Canal Zone (unpublished data from Panama Canal Company,
Engineering and Construction Bureau, Meteorological and
Hydrographic Branch).
Month
Station Rainfall in mm.
September I968
179.8
October
UtU.o
November
262.1
December
k6.2
January 1969
hh.2
February
13.2
March
10.9
8(4):133-152, 1969(1970) SEASONAL CHANGES
147
DISCUSSION AND CONCLUSIONS
There were far greater number of butterfly species and indi-
viduals active in the Clearing than in the Forest study areas,
both in the wet season and dry season. Part of the explanation
is likely an “overflow” effect, where many of the species that
normally fly high in the forest canopy come down low over the
clearing and are noted, but remain on top of the canopy and
thus unrecorded in the forest. Further reasons for the abund-
ance of species in the clearing undoubtedly lie in the helio-
thermic and thus heliophillic nature of butterfly physiology and
behavior (Emmel and Emmel 1962, 1963, 1964; Clench 1966;
Watt 1968). Most species, even in the tropics, require direct
sunlight to raise their body temperatures above ambient levels
for flight. The clearing also provides a much greater variety of
nectar sources for adult feeding, and a much greater variety of
of second-growth plants commonly used as larval foodplants in
such groups as the Pieridae, Nymphalidae and Heliconiidae.
1. Seasonal fluctuations in population size
There were considerable fluctuations in population size from
month to month for most species of butterflies on Barro Colorado
Island. These changes were usually associated with the change
from wet season to dry season, species flying mainly in one
season ( within the limits of the present survey ) . However, many
species reached their population peaks during the transition
period between wet and dry seasons. Fluctuations in populations
of papilionid and pierid butterflies are shown in Figure 2. These
changes may be due partly to changes in condition of larval
food, such as has been advanced as an explanaton of fluctuations
in tropical Drosophila populations (Pipkin 1953) where major
variations in population size follow variations in the local food
supply.
It is clear that later in the dry season, by the month of March,
the grasses and herbs of the Clearing area become very dry or if
still green, new growth has halted. Populations of some butter-
flies, such as the pierid Itaballia demophile, actually shift their
activity into the cooler more humid forest from the clearing
when the dry season is well underway (see Fig. 3). This shift
from open areas to the forest probably accounts for the increase
in number of species in the forest fauna in the dry season (Fig.
4, lower portion), although the dry season also probably pre-
sents more favorable environmental conditions for adult flight
148
EMMEL AND LECK
/. Res. Lepid.
Table 5 . Comparison of species recorded in the butterfly fauna
of Barro Colorado Island by Huntington (1932) and those censused
in the present study.
FAMILY Group Huntington Present Study
Papilionidae
5 species
6 !
Pieridae
13
10
Danaidae
U
2
Ithomiidae
11
6
Satyridae
l6
10
Brassolidae
2
1+
Morphidae
2
3
Heliconiidae
12
13
Nymphalidae
27
21
Riodinidae
1+0
11
Lycaenidae
3I+
U*
Hesperiidae
99
2*
* = only these species censused; others observed.
8(4):133-152, 1969(1970) SEASONAL CHANGES
149
and reproductive activities for the permanent forest species.
As just suggested, the influence of rainfall on adult activity
may also play an important role in causing seasonal population
fluctuations. The total number of hours of sun per day available
to the butterflies, for flying and reproductive activity was con-
siderably less in the wet season than in the dry season due to
afternoon cloudiness and rain. In a long-term or seasonal sense,
then, it is selectively advantageous to have a species' main flight
period in a time other than the wettest part of the rainy season.
The most advantageous time to fly and reproduce during the year
would seem to be the period immediately following the close
of the wet season, for later in the dry season (when environ-
mental conditions are still excellent for adult activity) the larval
foodplants may not be in suitable condition for feeding by
newly-hatched larvae. The apparent reality of this supposition is
reflected in the following data on changes in faunal organization
from the wet season to the dry season.
2. Seasonal fluctuations in species diversity.
When the number of species flying in the Clearing and Forest
study areas are graphed for each month (Fig. 4), it is clear that
(1) diversity in the Forest area increases in the dry season but
it is still at a relatively low level compared to that of the Clear-
ing fauna, and (2) diversity in the Clearing fauna, containing
clearly the species requiring a higher level of sunlight for activ-
ity, reaches a maximum diversity during the Transition Period
immediately following the Wet Season, before the Dry Season
conditions fully prevail.
This surprising confirmation of the preceding suppositions
(section 1) leads us to propose this as an example of a perhaps
more widespread phenomenon in the tropics: a “Seasonal Eco-
tone.” An ecotone, simply defined, is merely “a transition area
between two adjcent communities" (Webster’s New Collegiate
Dictionary). Treating the wet-season butterfly fauna and the
dry-season fauna as separate communities, the transition period
between the wet and dry seasons may be called a “Seasonal
Ecotone," and is simply a temporal analogy of the spatial con-
cept of an ecotone. This seasonal ecotone may be a general
phenomenon in influencing tropical species diversity, in that
one could find the greatest number of active species (of short-
life-cycle animals) between two distinct seasons, merely be-
cause both wet and dry season communities may be repre-
sented. The broader application of the seasonal-ecotone con-
cept is currently being considered for a number of tropical and
55
50
45
40
35
30
15
10
5
EMMEL AND DECK
J. Res. Lepid.
butterfly species observed in Clearing ( top ) and
7 areas per month from wet to dry season, 1968-1969.
8(4):133-152, 1969(1970) SEASONAL CHANGES
151
temperate animal groups (Emmel, in preparation). However,
it is clear with diurnally-active insects such as the butterflies
that the most reproductively favorable conditions also may
exist at this time and hence the seasonal ecotone fauna does
not merely represent an overlapping of communities but one
which has responded in an evolutionary sense to the most
satisfactory breeding period during the annual cycle (which
exists even in a tropical evergreen forest.) Preliminary review
of data from Costa Rican sites and elsewhere (Emmel, in
preparation) indicates that diversity increases only at the grad-
ual wet season-dry season seasonal ecotone (December in the
northern Neotropics), not at the sharp point of dry season-wet
season transition (April or May in the northern Neotropics).
This presumably is the result of dry-season-species’ adult in-
tolerance of the rainy conditions suddenly initiated by the start
of the wet season.
SUMMARY
Butterfly faunal censuses were made in a large clearing and
in the rain forest on Barro Colorado Island, Panama, during
the wet season and dry season, 1968-69. There were significant
changes in both population densities and species composition
(as represented by flying adults) from month to month and
between climatic seasons at this tropical site. These fluctu-
ations are apparently associated with available sunlight for
thermoregulation and with condition of larval hosts.
The greatest number of species flies at the transition period
between the wet and dry seasons. This ‘seasonal ecotone” is
probably due to both an overlapping of dry- and wet-season
faunas and to the favorable junction of environmental factors
for adult activity by tropical butterflies at that particular time.
ACKNOWLEDGMENTS
The Smithsonian Institution provided facilities and support
for this field study on Barro Colorado Island. The second-
named author was also supported by a Schuyler-Gage Fellow-
ship (Cornell University), and transportation was funded
through a Sigma-Xi Grant-in-aid-of-Research.
We thank Mr. Gordon Small, Canal Zone, for making many
of the species determinations of the B. C. I. butterflies.
LITERATURE CITED
ALCALA, A. C. 1966. Populations of three tropical lizards on Negros
Island, Philippines. Stanford University, California. Unpublished
Ph.D. dissertation.
152
EMMEL AND LECK
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ALLEE, W. C. 1926a. Measurement of environmental factors in the
tropical rain-forest of Panama. Ecology 7 1 273-302.
. 1926b. Distribution of animals in a tropical rain-forest with
relation to environmental factors. Ecology 7 : 445-468.
BATES, M. 1945. Observations on climate and seasonal distribution of
mosquitoes in eastern Colombia. J. Animal Ecol. 14: 17-25.
BOUGHEY, A. S. 1968. Ecology of populations. MacMillian Company,
New York. 135pp.
CLENCH, H. K. 1966. Behavioral thermoregulation in butterflies. Ecol-
ogy 47: 1022-1034.
EHRLICH, P. R., and P. H. RAVEN. 1964. Butterflies and plants: a
study in coevolution. Evolution 18: 586-608.
EMMEL, T. C., and J. F. EMMEL. 1962. Ecological studies of Rho-
palocera in a High Sierran community — Donner Pass, California.
I. Butterfly associations and distributional factors. Journ. Lepid.
Soc. 16: 23-44.
EMMEL, T. C., and J. F. EMMEL. 1963. Ecological studies of Rho-
palocera in a High Sierran community — Donner Pass, California.
II. Meterologic influence on flight activity. Journ. Lepid. Soc.. 17:
7-20.
EMMEL, T. C. 1964. The ecology and distribution of butterflies in a
montane community near Florissant, Colorado. Amer. Midi. Nat.
72: 358-373.
HIRTH, H. F. 1963. The ecology of two lizards on a tropical beach.
Ecol. Monographs 33: 83-112.
HUNTINGTON, E. I, 1932. A list of the Rhopalocera of Barro Colorado
Island, Canal Zone, Panama. Bull. Amer. Mus. Nat. Hist. 63: 191-
230.
JANZEN, D. H. 1967. Synchronization of sexual reproduction of trees
within the dry season in Central America. Evolution 21: 620-637.
JANZEN, D. H., and T. W. SCHOENER. 1968. Differences in insect
abundance and diversity between wetter and drier sites during a
tropical dry season. Ecology 49: 96-110.
LECK, C. F. 1970. Feeding behavior and ecology of fruit and nectar
eating birds in lower Middle America. Unpub. Ph.D. dissertation
(Cornell Univ. ).
MILLER, A. H. 1954. Breeding cycles in a constant equatorial environ-
ment in Colombia, South America. Acta XI Cong. Internat. Ornith.
Basel (1954): 495-503.
MOREAU, R. E. 1950. The breeding seasons of African Birds. I: Land
birds. Ibis 92: 223-267.
MOYNIHAN, M. H. 1968. Smithsonian Tropical Research Institute: 1967
among species of birds. Evolution 20:' 235-242.
PIPKIN, SARAH B. 1953. Fluctuations in Drosophila populations in a
tropical area. Amer. Nat. 87: 317-322.
PIPKIN, S. B., R. L. RODRIGUEZ, and J. LEoN. 1966. Plant host spe-
cificity among flower-feeding neotropical Drosophila (Diptera: Dro-
sophilidae). Amer. Nat. 100: 135-156.
RICKLEFS, ROBERT E. 1966. The temporal component of diversity
species of birds. Evolution 20: 235-242.
SEITZ, A. 1913. Macrolepidoptera of the world. Vol. 5. The American
Rhopalocera. Stuttgart.
SEXTON, O. J. 1967. Population changes in a tropical lizard Anolis
limifrons on Barro Colorado Island, Panama Canal Zone. Copeia
1967: 219-222.
SKUTCH, A. F. 1950. The nesting seasons of Central American birds in
relation to climate and food supply. Ibis 92: 185-222.
WATT, W. B. 1968. Adaptive significance of pigment polymorphisms in
Colias butterflies. I. Variation of melanin pigment in relation to
thermoregulation. Evolution 22: 437-458.
Journal of Research on the Lepidoptera
8(4):153-168, 1969(1970)
1160 W. Orange Grove Ave.^ Arcadia, California, U.S.A. 91006
© Copyright 1969
STUDIES ON NE ARCTIC EUCHLOE.
PART 6. SYSTEMATICS OF ADULTS
PAUL A. OPLER
Department of Entomology
University of California
Berkeley, California
This paper presents a discussion of some external morpho-
logical features of the adults of Nearctic Euchloe, a key to
identification, descriptions of named entities, and illustrations.
Stress will be given those features found most useful in the
separation of adults.
METHODS
CHROMATOGRAPHY. The procedures followed were based
on those of Hadorn and Mitchell (1951) and Biserte (1960).
Solvent systems of n-Propanol and aqueous ammonia (2:1) and
Butanol, glacial Acetic acid, and water (4:1:5) were employed.
The chromatograms were obtained by a uni-directional ascend-
ing method and the spots were revealed under ultra-violet
illumination.
EXTERNAL FEATURES. With the exception of androconial
scales, all external features were studied with the aid of a
dissecting microscope at 10, 30, or 60 power.
Measurements of wing length were made with a vernier
caliper to the nearest one-tenth millimeter. Measurement of
costal length was made from the point of wing attachment to
furthest extent of the apex, not including the fringe.
The width of the black bar at the end of the cell on the dorsal
surface of the forewing was measured by counting the number of
scale rows on the right wing under 30 power from the first row
with 50% or more black scales to the first comparable row on
the opposite side. The number of white scales in the bar were
153
154
PAUL A. OPLER
J. Res. Lepid.
Fig. 1. Upper Row. Left: Euchloe creusa, male, upper surface; Right;
Euchloe creusa, male lower surface; Middle Row. Left: Euchloe olympia,
male, lower surface; Lower Row. Left: E. ausonides, male, upper surface;
Left: E. ausonides, male, lower surface.
H(4):153-16H, 1969(1970) NEARCTIC EUCHLOE
155
counted below the costal vein, as the bar is frequently ill-defined
above this vein. Any white scale completely surrounded by
black scales was considered as occurring within the bar. For
individuals with more than fifty white scales in the bar, a por-
tion of the bar was counted for the character, and the total was
then arrived at by extrapolation.
The relative length of the radial veins were compared with
the aid of an ocular grid. If one does not clear the wings, the
veins are best observed on the ventral surface of the forewing
with light from the illuminator striking the wing at an oblique
angle.
Androconial scales were studied by scraping the area of the
bar on the dorsal surface of the forewing with an insect pin or
dissecting needle, transferring the scales to a microscope slide,
covering them with a cover slip, searching for the proper scales
under low power, and finally studying them under 200 to 400
power with a compound microscope. For permanent prepar-
ations, a mounting medium should be applied around the edge
of the cover slip only, and pressure applied to the cover slip until
the preparation dries.
GENITALIC PREPARATIONS. The genitalia were subjected
to the usual preparatory procedures but were not mounted on
slides. Genitalia were observed in a mixture of ethanol and
glycerine in a small dissecting dish and were stored in small
vials inside larger museum jars.
DRAWINGS. The subjects for the figures were observed
through a binocular microscope equipped with an ocular grid.
Pencil drawings were made on grid paper, and later the originals
were traced onto finer grade paper and inked in.
MORPHOLOGICAL FEATURES
PIGMENTATION . It is well known that a group of pigmental
compounds known as pterines is responsible for the white, yel-
low, and red wing colors of many members of the family Pieri-
dae. Since these compounds have been demonstrated to occur
in the wings of a species of Anthocaris by Good and Johnson
(1949), and since the Euchloe possess white and yellow wing
pigments, I decided to demonstrate the presence of pterines in
the wings of Euchloe.^ Specimens of Euchloe ausonides and
E. hyantis lotta, as well as other species of Pieridae, were
used in the experiment. Light blue fluorescent spots with RF
1 Chromatography experiment conducted in insect physiology laboratory at San Jose
State College, Dr. Ballard, instructor.
156
PAUL A. OPLER
/. Res. Lepid.
Fig. 2. Upper Row. Left; Euchloe hy antis, male, upper surface; Right:
E. hyantis, male, lower surface; Middle Row. Left; E. creusa, male, right
forewing; Right: E. olympia, male, right forewing; Lower Row. Left: E.
ausonicles, male, right forewing; Right: E. hyantis, male, right forewing.
8(4):153-168, 1969(1970) NEARCTIC EUCHLOE
157
values of 0.21 were obtained for both species of Euchloe with
the propenol-ammonia solvent system. Since this finding was
also produced with a wing sample of Pieris rapae L. it was
tentatively assumed that leucopterin, the pigment responsible
for the white wing color of many Pieridae, was the compound
which formed these spots. Light blue fluorescent spots with RF
values of 0.35 and barely discernable purple fluorescent spots
with RF values of 0.29 were obtained for both species with
the butanol-acetate- water solvent system. It was deduced that
these values possibly represented breakdown products of xantho-
pterin, the pigment responsible for the yellow wing colors of
many Pieridae. Needless to say, these results are far from de-
finitive. It was realized that either larger samples or more re-
fined techniques should be used in conjunction with chemically
defined standards if significant differences are to be shown be-
tween species or populations of Euchloe.
The differences between species, populations, and individuals
with regard to the whitness of wing color may be due to the
presence of varying proportions of xanthopterin mixed with
leucopterin. The pearly lustre or sheen or its absence are best
explained by physical effects, i.e., the presence of ridges on the
scales, the angle of scale elevation from the point of attachment,
thickness of scales.
SCALE TYPES. The ‘marbling” on the ventral surface of the
hindwings is composed of two types of scales. The first type,
which consists of the white and yellow scales, is of roughly rec-
tangular outline with lobes or teeth on the distal margin. There
appear to be differences in the number and outline of the lobes
or teeth between different populations or entities of Euchloe.
However, a satisfactory method of noting these differences was
not arrived at in the course of this study. The black scales on
the ventral surface of the hindwing, which together with the
yellow scales give the visual effect of green “marbling”, are
ovoid in outline and are dentate on the distal margin with the
exception of many individuals of Euchloe ausonides coloradensis.
The distal margin of the black scales of these individuals is
simple.
The males possess androconial scales on the dorsal surface
of the forewings in the area of the black bar marking located
at the distal margin of the discal cell. The location of these
TABLE 1 . STATISTICAL SURVEY OF SOME WING CHARACTERS
158
PAUL A. OPLER
/. Res. Lepid,
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8(4):153-168, 1969(1970) NEARCTIC EUCHLOE
159
scales has never been reported for members of the tribe
Euchloini. Warren (1961) and Chang (1963) have reported
that the androconial scales have a distinctive shape which is
constant for any given species of the genus Pieris. It was hoped
that these scales would furnish similar diacritical differences iA
the Nearctic Euchloe, but upon microscopic examination they
were found to show very slight interspecific differences. While
the androconial scales of the Ausonides species group were
relatively constant in having the lateral edges of the scales
approximately parallel or slightly divergent, the androconial
scales of the Hyantis complex were found to be quite variable.
On more than one occasion scales varying from ovoid to trape-
zoidal were found on one specimen of Euchloe hyantis loUa.
BAR CHARACTERS. Since Brown (1955) reported that
Euchloe hyantis lotta can be distinguished from E. ausonides
coloradensis by its wider bar marking, it was decided early in
the study to use this as a possible character in the study. The
bar marking was measured by counting its width at a point near
the middle in scale rows, i.e. the number of scales encountered
in a line across the marking. All specimens recorded in the study
were coded for this character. It was found that although indi-
vidual variation was wide it did give a good measure of
difference between certain entities (see Table 1).
In looking at specimens with intent to code for the above
character, it was discovered that all specimens of E. ausonides
possessed a scattering of white scales within the bar, while
inividuals of E. hyantis did not. Hence, the writer coded all
specimens for the number of white scales in the bar marking.
This character appears to be the best qualitative means of sep-
arating adults of Euchloe ausonides from Euchloe hyantis with-
out resorting to dissection of the genitalia. Occasionally worn
individuals of E. ausonides, especially females, will not display
this character well as the scales appear to be more deciduous
with age than are the other scales on the wings. Some popula-
tions of Euchloe hyantis that occur in the middle elevations of
the Sierra Nevada of California are composed of individuals
which so closely resemble E. ausonides from nearby areas that
only by examining the individuals in question under a binocular
microscope for this character can one be sure which species
he is dealing with. The genitalia of such individuals subse-
quently support the conclusions which were arrived at on the
basis of the presence or absence of white scales in the bar
160
PAUL A. OPLER
J. Res. Lepid.
marking. When other evidence, which will be presented in later
papers of this series, indicates that Euchloe hy antis must have
become isolated from the line which gave rise to the radiation
of the Ausonides species group at a relatively early date, one
must realize that this is either a startling example of convergent
evolution or an improbable coincidence.
VENATION. In the past, several workers, including Dyar
(1894) and Grote (1900), have proposed that members of the
genus Euchloe can be discriminated on the basis of wing
venation, while other writers such as Butler (1899) and Klots
(1930a) have argued against the wisdom of employing this
character. The radial veins on the forewing were usually used
in attempts to utilize wing venation as a classificatory aid for
Euchloe. The antagonists to such hypotheses reasoned that
these characteristics were variable from one specimen to an-
other. The present writer found that although the state of the
radial veins varied slightly from one individual to another,
definite trends for each species were clearly discernible, (see
Fig. 4). The method employed was to contrast the length
of the stem of the fourth and fifth radial veins from the bifurca-
tion of the third radial with the length of the fourth radial vein.
A trend for Euchloe creusa could not be noted owing to the
small sample that was available, however specific characteristics
were found for the other three species. The fourth radial of
Euchloe olympia was invariably longer than its stem, the length
of the fourth radial of E. ausonides was shorter or about equal
to the length of its stem, and the fourth radial of individuals
of Euchloe hyantis was always shorter than its stem. In fact,
the fourth radial vein of both wings of many individuals of
E. hyantis was found to be absent or only barely present.
EXTERNAL GENITALIA. An excellent world-wide tribal
revision of the Euchloini by Klots ( 1930a ) was based in large
part on the structure of the external genitalia of the male
insects. In spite of that fact, no satisfactory genitalic character-
istics have been reported at the species level for any of the
Nearctic Euchloe. As with many other characteristics of this
subgenus, the external genitalia are perplexingly similar in
superficial appearance. As a result of the study reported in
this paper several features of the genitalia were found which
will readily separate individuals of the two species groups in-
volved. The outline of the juxta, when viewed from the pos-
terior angle, is V-shaped for individuals of Euchloe ausonides
8(4):153^168, 1969(1970) NEARCTIC EUCHLOE
161
and is Y-shaped for the other three species (see Fig, 3). For
species of the Ausonides group, the lateral edges of the tegumen,
when viewed from the dorsal aspect, are parallel and do not
converge until just prior to the point of juncture with the uncus,
while for individuals of the Hyantis complex, the lateral edges
of the tegumen are noticeably convergent distally or are irregular.
The saccus of members of the Ausonides group tends to be
regular in outline, while the saccus of individuals of the Hyantis
complex is irregular in outline. The cucullus of members of
the Hyantis complex terminates abruptly after the elaboration
of the distal tooth, while the cucullus area of the valvae of
members of the Ausonides group extends a short distance be-
yond the distal tooth (Fig. 3).
KEY TO THE ADULTS OF NEARCTIC EUCHLOE
1. Length of R4 much less than length of stem R4 5, white ground on
ventral surface of hindwing usually with pearly lustre, bar at end
of cell on dorsal surface of forewing containing less than five white
scales below costal vein, cucullus area of valvae terminating abruptly
after distal tooth, lateral margins of uncus becoming convergent
distally (dorsal view) Euchloe hyantis complex 7
Length of R4 about equal to or greater than length of stem R4 5,
white ground on ventral surface of hindwing usually dull white,
bar at end of cell on dorsal surface of forewing containing five or
more white scales below costal vein, cucullus area of valvae not
terminating abruptly after distal tooth, lateral margins of uncus
parallel or only slightly convergent distally Ausonides species
group 2
2(1). Juxta of male V-shaped, sterigma usually evenly curved in lateral
or ventral view, female sometimes with dorsal surface of hindwing
distinctly yellowish in color in comparison to ground of forewing .. 3
Juxta of male Y-shaped, sterigma sinuous in lateral of ventral view,
ground color of dorsal surface of hindwing almost never yellowish.. ..6
3(2). Bar at end of cell on dorsal surface of forewing narrow, black scales
on ventral surface of hindwing often ovoid in outline, a higher per-
centage of black scales as compared to yellow scales on ventral
surface of hindwing, southern Rocky Mountains of northern New
Mexico, Colorado, and southern Wyoming, often at high altitudes
Euchloe ausonides coloradensis
Bar at end of cell narrow or wide, black scales on ventral surface
of hindwing with two or more teeth or lobes distally, black scales
and yellow scales in about equal numbers on ventral surface of
hind wing, not occurring in the areas listed above 4
4(3). Bar narrow, hindwing of female barely contrasting in color with
forewing, low mountains of west-central Manitoba Euchloe
ausonides mayi
Bar wide, usually more than eleven scale rows in width, hindwing
ground of female usually contrasting with that of dorsal surface
of forewing, not occurring in west-central Manitoba 5
162
PAUL A. OPLER
J. Res. Lepid.
Fig. 3. Lateral view of male genitalia. A. E. creusa. B. E. olympia. C.
E. ausonides. D. E. hyantis. E-F. Posterior view of male genitalia showing
two configurations of juxta.
8(4):153~168, 1969(1970) NEARCTIC EUCHLOE
163
5(4). Female possessing one of three phenotypes, i.e. both wings white
dorsally, both yellow, or forewing white and hindwing yellowish,
occuring in the Coast Range, Sacramento Valley, and northern San
Joaquin Valley in California from Mendocino County south to
Monterey County Euchloe ausonides ausonides
Female with dorsal surface of hindwing always distinctly yellowish,
not in the lowlands of central California, southern Rocky Mountains,
or west-central Manitoba Euchloe ausonides ssp,
6(2). Antennae clothed with white scales only, marbling on ventral sur-
face of hindwing strongly reduced, black marking on apex of fore-
wing often reduced, buff-colored scaling usually not present on
costal margin of forewing, black scaling not invasive on dorsal
surface of hindwing, occurring in eastern half of United States and
adjacent portions of Canada in Manitoba and Ontario ........ Euchloe
olympia
Antennae clothed with both white and black scales, marbling on
ventral surface of hindwing often heavy and of a “broken’’ nature,
black marking on apex of forewing not reduced, buff-colored scaling
present on costal margin of forewing, black scaling at base of hind-
wing on dorsal surface invasive outwardly more so than other
species or Nearctic Euchloe, occurrence associated with mountain
cordillera of Canada and Alaska , occurring near timberline, i.e.
7000’ in southern Alberta, 4000’ in northern British Columbia and
close to sea level in Northwest Territories (McKenzie River
delta) Euchloe creusa
7(1). Occurrence associated with Northern Desert Scrub (sagebrush)
or Southern Desert Scrub Biomes ( desert ) west of the Continental
Divide (except Rio Arriba County, New Mexico) and east of the
Sierra Nevada and Cascade Mountains Euchloe hyantis lotta
Occurrence in the north Coast Range of California, west slope of
the Sierra Nevada, Transverse Ranges of southern California ( ex-
cluding lower desert slopes), and the Peninsular Ranges of San
Diego County, California and Baja California del Norte 8
8(7). Occurring in the north Coast Range of California from Sonoma
County north to Siskiyou County Euchloe hyantis hyantis
Not occurring in the north Coast Range of California 9
9(8). Occurring in the higher portions of the San Bernardino Moun-
tains Euchloe hyantis andrewsi
Not occurring in the higher portions of the San Bernardino Moun-
tains of southern California Euchloe hyantis ssp.
Euchloe ausonides ( Lucas )
Male. — Forewing length, 21 mm. Antennae: brownish-tan,
outer surface of shaft clothed with black and white scales, black
predominating, nudum and inner surface of shaft naked, tip of
nudum with small microtrichia; labial palpi twice as long as
head, directed anteriorally at a slight dorsal angle, clothed with
black and white elongate scales, about three and a half times
as long as wide, long white hair-like scales on inner face, similar
black scales directed ventrally, a group of longer scales, both
white and black, projecting from ventral base of palpi; head
164
PAUL A. OPLER
J. Res. Lepid
Fig. 4. Venational configuration of radial group of right forewing. A. E
ausonides. B. E. creusa. C. E. olympia. D. E. hyantis.
8(4):153-168, 1969(1970) NEARCTIC EUCHLOE
165
black with eyes green, frons with prominent tuft of long black,
white, and yellowish hair-like scales, primarily white ventrally
and yellowish laterally; vertex with long white hair-like scales, a
patch of yellow and black hair-like scales half the length of
those on center of vertex present on lateral margins of vertex,
a group of shorter bright yellow scales between eyes and base
of antennae; eyes bordered dorso-posteriorally by bright yellow
and black flattened scales, a collar of bright yellow hair-like
scales on cervical region adjacent to posterior and ventral margin
of eyes.
Thorax: clothed with black appressed quadrate flattened scales
and long hair-like scales, whitish-gray throughout 9/10 of length
and black at base; pleuron covered with yellow sub-elliptic
flattened scales and long yellowish hair-like scales; legs with
femora covered with white flattened scales becoming tan distally,
also with long white hair-like scales predominately on ventral
surface and becoming shorter distally; tibia, tarsi and pretarsi
brownish-tan, covered with stout setae, narrow white flattened
scales on sparsely clothed tibia and tarsis. Wings: forewing with
costal and outer margins slightly curved, inner margin straight,
outer margin pointing outward anteriorally giving wings a sightly
pointed look, stem R4 5 longer than Rs, upper surface completely
clothed with flattened dull-white scales in approximate vertical
rows except as follows: black flattened scales occurring solidly
on basal one-tenth of wing, on costal margin of wing as eight
small vertical marks extending to cell, on apex in typical Euchloe
manner with intermixed white scales from Ri to Ms at distal ends
of veins, and at distal end of discal cell as patch about thirteen
scale rows wide with about one hundred white scales intermixed,
long grayish-white hair-like scales coinciding with basal patch
of black scales, yellow-buff narrow flattened scales extending
along costal margin from base to apex, fringes (along inner and
outer margins) composed of long white hair-like scales, black
hair-like scales on fringe at terminus of Rs, Mi, M2, Ms, and Cui.
Hindwing above with dull white scales as on forewing, black
flattened scales on basal area of wing, extending outwardly
further than on forewing, at stem of cubitus, and at terminus
of all veins coinciding with long black hair-like scales on fringe;
ventral surface of forewing with white scales as above, black
scales as above on costal margin and outer margin, black patch
at end of discal cell not as extensive as on upper surface and
white scales absent, black scales absent at base of wing and
166
PAUL A. OPLER
J. Res. Lepid,
much less extensive on apical area, yellow-bufiF scales as on
upper surface, sparsely distributed white hair-like scales occur-
ring anterior to cubitus and extending to outer end of discal cell,
flattened yellow scales occurring with black scales on apical
area giving greenish appearance; lower surface of hindwing with
flattened slightly dentate white scales in rough rows in between
complex “green” marbled pattern produced by intermixing of
flattened black and yellow scales, long hair-like scales, white on
white areas and pale yellow over marbling extending from base
of wing approximately to an imaginary line from distal end of
anal margin to distal end of inner margin, a small patch of
flattened black scales contiguous with marbling at Mu-Cu with
one long black hair-like scale. Abdomen: dorsum clothed with
flattened black scales intermixed with a few flattened white
scales, white scales increasing and black scales decreasing ven-
trally until all white on sternum, long grayish hair-like scales
on anterior half of abdomen and along entire length on sternum,
white slightly spatulate scales sparsely covering posterior half
of abdomen and densely covering posterior margin of segment
eight and outer surface of valvae.
Female. — Forewing length, 22 mm. As in male except patch
at end of discal cell of forewing about eighteen scale rows in
width with about twenty white scales intermixed; hindwing
above with scales yellow-cream in color; scales on lower surface
of forewing largely buff in color, about fifty white scales in
center of patch at end of FW discal cell ventrally.
Euchloe creusa (Doubleday)
Male. — Forewing length, 18 mm. Antennae brownish-tan,
outer surface clothed with black and white scales, white pre-
dominating; hair-like scales on dorsal surface of thorax as in
E. ausonides but denser, yellow flattened and long yellow hair-
like scales on pleuron; legs with long black and white hair-like
scales primarily on ventral surface of femora, white predominat-
ing; forewing with stem R4 5 about equal in length to Rs, upper
surface of forewing with eight black marks on costal margin,
“Euchloe” mark at apex with white area above Mi not as a well-
defined circle, instead the effect is of a diagonal bar beginning
between Ra and R4 and ending between M2 and Ma; bar mark
at distal end of cell about eight scale rows in width with about
35 white scales intermixed; flattened black scales extending into
cell from basal area on dorsal surface of hindwing; white scales
8(4):153^168, 1969(1970) NEARCTIC EUCHLOE
167
on ventral surface of hindwing more iridescent than those of
E. ausonides; marbling more extensive and irregular than that
of other Nearctic species. Abdomen: flattened black scales on
dorsal and pleural areas with only an occasional white scale;
venter covered with a mixture of white and pale yellow flattened
scales; long hair-like scales covering entire surface of abdomen,
gray on dorsal and pleural areas, yellowish ventrally.
Female. — Forewing length, 17.8 mm. As in male except patch
at end of discal cell on forewing about sixteen scale rows in
width with about seven white scales intermixed; flattened black
scales on upper surface of hindwing as in male but some present
on all areas of wing; a small patch of about 30 black scales
present on Idc; about 25 dull gray scales in center of patch at
end of discal cell on ventral surface of forewing.
Euchloe olympia (Edwards)
Male. — Forewing length, 18.5 mm. Antennae: outer surface
of shaft and most of club clothed with small white flattened
scales; labial palpi about one and a half times as long as head,
clothed with white elongate scales, about four times as long as
wide, long white hair-like scales projecting downward and in-
ward, a few long black hair-like scales intermixed on outer face;
frons with prominent tuft of long white and black hair-like
scales directed anterad slightly beyond tips of palpi, white
mesially with some black scales laterally; pleuron covered with
yellow sub-elliptic flattened scales and long yellow hair-like
scales; legs with scaling as for E. ausonides. Wings: Rs almost
twice length of stem R4 5, upper surface with white ground
slightly more iridescent than that of E. ausonides; black macula-
tion on apex reduced to three small patches, one just basal to
R3 bifurcation, one at distal end of Ma, and one composed of
scattered black scales near R4 5; four small vertical marks in
C-Sc formed by small patches of black scales; patch at distal
end of discal cell on dorsal surface of forewing about 13 scale
rows in width with about five white scales intermixed; black
scales not present in fringe; buff scales absent from costal
area; hindwing with long black scales in fringe at termini of Rs
and Mi; ventral surface of forewing with black scales as above
patch at R4 5 absent, flattened yellow scales occurring with black
scales in two apical patches giving greenish appearance; patch at
end of discal cell much less extensive than on dorsal surface with
about 100 white scales in central portion; ventral surface of hind-
168
PAUL A. OPLER
J. Res. Lepid.
wing with marbling pattern strongly reduced. Abdomen: dorsal
and pleural areas clothed with flattened black scales with a few
white scales, venter clothed with white scales.
Female. — Forewing length, 19.2 mm. As in male except patch
at end of discal cell on forewing about 14 scale rows in width
with no white scales intermixed; about 50 white scales in central
portion of patch at distal end of cell on ventral surface of fore-
wing.
Euchloe hy antis (Edwards)
Male. — Forewing length, 17.5 mm. Outer surface of antennal
shaft and club clothed with white and black scales, white pre-
dominating; labial palpi lacking black elongate scales as in E.
ausonides, frons lacking long yellowish hair-like scales; patch of
hair-like scales at lateral margin of vertex with white and black
scales; a group of short pure white scales between eyes and base
of antenna; on forewing stem R4 5 much longer than Rs; patch
at distal end of discal cell on dorsal surface of forewing about
13 scale rows in width with only two white scales intermixed;
buff scales absent from costal margin; ventral surface of forewing
with patch at distal end of cell about as extensive as on dorsal
surface with no perceptibly lighter scales in center.
Female. — Forewing length, 18.1 mm. As in male except patch
at distal end of cell on dorsal surface of forewing about 16 scale
rows in width with no white scales intermixed; about 70 light
gray scales in center of patch at end of FW discal cell ventrally.
ADDENDUM
After the manuscript for this paper was submitted for publi-
cation, the materal of this genus contained in the Canadian
National Collection was examined. Since the material there
included important distributional additions, the data for their
material from Alaska and Canada are presented below. It should
be noted that none of this information has been incorporated
on the distribution maps.
Journal of Research on the Lepidoptera
8(4):169-176, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
LABORATORY PRODUCTION OF THE
MONARCH BUTTERLY, Danaus plexippus
FRANCIS MUNGER and THOMAS T. HARRISS
Department of Biology, Whittier College, Whittier, California
There is need for monarch butterflies, Danaus plexippus
(Linn.), for scientific and educational work. For these purposes
equipment and methods for rearing the butterfly are being
developed.
In the feld, in regions where the monarch occurs, in season,
the butterfly is attracted to milkweeds for egg-laying. Plantings
of Asclepias curassavica are especially useful for luring migrating
butterflies to obtain them for experimental work. This plant is
perennial and sub-tropical. In addition to milkweed flowers, the
butterflies are attracted to many other kinds of flowers for their
nectar.
In the laboratory the butterflies will drink from damp paper,
water, and water to which honey has been added. Honey was
added to the water at the rate of 1 teaspoon per cup of water.
Crumpled paper toweling of a stiff variety or newspaper was
placed in a shallow dish containing the solution. The butterflies
would stand on, and drink from the damp paper.
Oviposition cage. An essential tool for rearing the butterfly is
the oviposition cage. The cage used by the authors (Fig. 1) is
17 inches tall, 2014 inches wide, and 12 inches deep. The bottom
is Yz inch plywood to which a wooden frame is attached. One
side is provided with a terry cloth sleeve 6 inches in diameter
and 10 inches long attached to wooden panel. Through this
sleeve, butterflies, glass tumblers with water, cuttings of milk-
weed plants, and cut flowers can be passed without danger of
butterflies escaping. The top, back, and one side of the cage
are covered with muslin. The cloth at the back is in the form of
a curtain, fastened at the top and weighted at the bottom with
a piece of masonite 14 by 3 by 20 inches. The masonite
is held against the back of the cage by sheet-metal guides. The
front of the cage is covered with glass or plexiglas.
Present address of Thomas T. Harriss: Department of Biology, Western
Colorado State College, Gunnison, Colorado 81230.
169
170
MONGER AND HARRISS
/. Res. Lepid.
Fig. 1. — Oviposition cage.
8(4}:169-176, 1969(1970)
PRODUCTION OF MONARCH
171
When in operation, the cage contains a 9 by 1 Yz inch aluminum
cake pan with honey-water and crumbled paper, a glass tumbler
containing water and cuttings of milkweed, A. currassavica, and
a number of egg-laying females.
Under the artificial conditions of the oviposition cage the
butterflies will not always mate when they become mature. It is
usually necessary to force-mate the butterflies by a method used
by the workers at the University of Toronto, Canada. ( Urquhart
1965).
Incubation of eggs. Eggs are laid almost exclusively on the
leaves and stems of milkweed. Occasionally some eggs are laid
on the damp paper, on the tumbler, or on the wood frame of
the cage. The milkweeds are replaced with fresh plants every
1 to 3 days. Small parts of leaves and stems bearing 1 or several
eggs are cut from the plants, placed in a pile on a piece of glass
AYz inches square, and covered for incubation with a clear, plastic
cup SYi inches in diameter and 3 inches tall (Fig. 2). A dispos-
able cup of this size is obtainable from most retail liquor stores.
Any number of eggs up to 100 may be incubated at one time
under a cup. Humidity must be kept low enough to prevent the
growth of mold, which seems to kill the eggs.
Rearing the caterpillars. When the eggs hatch, most of the
larvae crawl up on the sides of the cup, now a cage, and rest for
a time before they are ready to feed. At this time 1 or 2 milkweed
terminals composed of 4 to 6 leaves each are placed in the cage,
partly in contact with the plastic surface. Before long the
caterpillars transfer to the milkweed and commence to feed. As
the caterpillars grow and become crowded they are distributed
among other cages. Three fifth instar caterpillars can be
reared to maturity in 1 cage if they are of slightly different ages
so that they do not interfere with each other when they are
preparing to suspend themselves.
Storage of pupae. Pupae can be held in the cage until the
butterflies emerge, or they can be removed and stored on a
string rack. This rack is a wood frame with strings stretched
horizontally 4 inches apart. A cardboard try beneath the
strings is used to collect tachinid parasites issuing from
suspended insects. Parasites may be present in larvae collected
in the field, but not in laboratory-reared material. .
The silk to which the pupae are attached will peel off the
plastic surface if it is first started by rubbing with the finger or a
rubber eraser. A mounting device is made of a 114 -inch long
piece of %-mch masking tape. First, fold the masking tape at
172
MUNGER AND HARRISS
}. Res. Lepid.
Fig. 2. — Plastic cup cage for incubating eggs and rearing small caterpillars.
Fig. 3. — Cardboard cylinder rearing cage.
8(4):169~176, 1969(1970)
PRODUCTION OF MONARCH
173
one end, lengthwise, half way. Then, place the silk of the pupa
on the other end and press the adhesive surfaces together with
the silk between. Th pupa can then be fastened to the string by
a slanting cut in the folded tape or by folding i/4-inch of
the tape over the string and securing it with a small paper
clip. The pupa can then be identified by marking a number on
the masking tape.
Cardboard rearing cage: A cage for rearing a larger number
of fifth instar caterpillars to maturity per cage is composed of
a cardboard cylinder 6 inches in diameter and 3 inches tall
(Fig. 3). The top is a 7-mch square piece of glass or plexiglas. A
7“inch square piece of paper is laid on the bottom glass to absorb
moisture and to keep the glass clean. The capacity of this cage is
about 15 insects. When the caterpillars finish feeding, they
crawl to the cover, form the silk button, and suspend themselves.
Pupae suspended in this manner can be stored in a slotted
wooden rack (Fig. 4). If desired, the pupae can be removed
from the glass cover, either by scraping the silk off with a razor
blade, or peeling it off wet. The insects can then be fastened to
masking tape as described earlier. The removal of pupae from
plexiglas is simpler. The silk peels off easily, dry.
It is more efficient to rear the larger number of caterpillars
at one time, particularly if there is no virus disease (Urquhart,
1966 and Urquhart and Stegner, 1966) present. An advantage in
rearing the smaller number of caterpillars in one cage is that if
one insect is infected with the virus disease common to the mon-
arch butterfuly, only 2 additional specimens are exposed. Also,
the plastic cages are more easily sterilized without damage to
them.
The caterpillars were reared in a dry, well-ventilated basement
room with daylight, at about 74 degrees F. Under conditions
of high humidity the silk may not adhere well to the plastic,
but it never fails to stick to the glass.
Egg-production. An egg-production experiment was carried
out in the Whittier College greenhouse from May 15 to June 27,
1968. The object of the experiment was to determine the egg-
laying potential and longevity of butterflies which were fed on
honey-water and flowers, and on honey-water alone. The fre-
quency with which the butterflies feed on flowers in the field
suggests that flowers might be necessary for the greatest egg-
production in a cage.
174
MUNGER AND HARRISS
J. Res. Lepid.
Fig. 4. — Wooden rack for storing and displaying suspended pupae.
The greenhouse is partially shaded by eucalyptus trees. The
top ventilator was open continuously. Maximum and minimum
temperatures F. were read on 26 and 25 days, respectively of
the 43 days of the experiment. The maximum, minimum, and
average temperatures for the maximum temperatures were 95, 74,
and 82.2 degrees. The maximum, minimum, and average tem-
peratures of the minimum temperatures were 67, 54, and 56.8
degrees.
The butterflies used in the experiment were reared from eggs
which had been laid by a number of laboratory-reared butterflies.
8(4):169-176, 1969(1970) PRODUCTION OF MONARCH 175
TABLE 1 . DATES BUTTERFLIES WERE MATED AND INTRODUCED
INTO OVIPOSITION TEST CAGES.
Date transferred to cage
Butterfly
Date
Date
Honey-water
Honey-water
number
emerged
mated
and flowers
alone
1
May 10
May 16
May 15
2
May 10
May 16
May 15
3
May 17
May 22
May 22
4
May 17
May 22
May 22
5
May 18
May 23
May 23
6
May 18
May 23
May 23
7
May 19
May 25
May 25
8
May 19
May 25
May 25
TABLE 2.
LONGEVITY AND
EGG-PRODUCTION OF 8
BUTTERFLIES, 4
FED ON
HONEY- WATER AND FLOWERS, AND 4 FED ON HONEY-WATER ALONE.
CONDITION
Honey-water and flowers Honey- water alone
Longevity
Total eggs
Longevity
Total eggs
Butterfly
Days
Butterfly
Days
1
41
2
34
3
37
4
40
5
36
6
44
7
35
1551
8
44
2365
re.
37*1
387.8
40.5
591.3
176
MONGER AND HARRISS
/. Res. Lepid.
Procedure. Two oviposition cages were used with 4 force-
mated butterflies distributed to each cage as shown in Table 1.
Each cage contained honey-water and milkweed leaves. One
cage contained, in addition, a separate tumbler with fresh-cut
flowers— scabiosa, lantana, orange, milkweed, and other flowers.
The flowers were attractive to the butterflies, and they were
observed to feed on them frequently. The milkweed leaves,
upon which the eggs were laid, were replaced with fresh leaves
at from 1 to 3-day intervals, and the eggs counted. Tagged
butterflies were used in the experiment. The longevity of each
butterfly, therefore, was measurable, but it was not possible to
determine the egg-production of the individual butterfly.
Results. The results of the experiment are shown in Table 2.
The butterflies in the cage without the flowers laid 60.8 percent
of all the eggs, an average of 591.3 eggs per butterfly. These
butterflies lived an average of 40.5 days, 3.4 days longer than
those with the flowers.
It is clear, contrary to what might be expected, that the
flowers added nothing to the egg-laying ability of the butter-
flies. On the contrary, the flowers seemed to detract from the
capacity of the butterflies to lay. A possible explanation of this
result, suggested by Dr. Hovanitz, may be that time spent on
the milkweed leaves may have been reduced by the attraction of
the flowers. From a practical standpoint, it is convenient that
flowers do not seem to be an advantage in egg-production.
The number of eggs laid by the butterflies with flowers was
about equal to what Urquhart (1960) suggested might be ex-
pected to be laid under ideal conditions. He examined monarch
ovaries and found more than 400 eggs.
Note: Seeds of Asclepias curassavica are available from Clyde
Robin, P. O, Box 2091, Castro Valley, California, and from
Pearce Seeds and Plants, Moorestown, New Jersey 08057.
REFERENCES
URQUHART, F. A., (1960). The Monarch Butterfly. The University of
Toronto Press, Canada.
(1965). Personal communication.
( 1966 ) . Virus-caused epizootic as a factor in population fluctu-
ations of the Monarch butterfly. ]. Invert. Path. 8:492-495.
and R. W. STEGNER, (1966). Laboratory Techniques for Main-
taining Cultures of the Monarch Butterfly. Jour. Res. Lepid. 5(3): 129-
136.
Journal of Research on the Lepidoptera
8(4):177-181, 1969(1970)
1160 W. O'^ange Grave Ave-, Arcadia, California, U.S.A. 91006
© Copyright 1969
OBSERVATIONS AND NOTES ON
THE REARING OF PAPILIO INDRA
KAIBABENSIS ( PAPILIONIDAE )
RONALD S. WIELGUS
3434 W. Augusta, Phoenix, Arizona 85021
On 31 May 1970 five larvae of Papilio indra kaibabemis
Bauer were found by the author on individual host plants of
Pteryxia petraea (Jones) Coult. & Rose growing on the slopes
and along the North Kaibab Trail opposite Roaring Springs,
North Rim, Grand Canyon National Park, Arizona. These were
taken back to Phoenix on 1 June 1970 along with a small amount
of host plant material in an attempt at rearing. No previous at-
tempt at rearing the larvae of this choice swallowtail had been
made by the author.
The larvae found ranged as follows: one first-instar, one
second-instar and three third-instars. These were numbered
1 through 5 respectively, to facilitate recording of individual
behavior during the rearing process. Upon arrival at Phoenix it
was discovered that larva No. 5 had moulted sometime during
the several hours’ return drive.
The larvae were kept indoors at a constant 80° F. and fed on
the leaves of the host plant for five days, to 6 June 1970, during
which time the host plant, in a vase of water, dehydrated and
became stiff and brittle. By this time four of the five larvae had
moulted and there were now one first-instar, one third-instar,
two fourth-instars and one fifth-instar. It was immediately ap-
parent that successful continuation of the rearing was dependent
upon the acceptance by the larvae of substitute host plant.
Emmel and Emmel (1967) found Tauschia arguta (T. & G. ) to
be an acceptable substitute and successfully reared kaibabemis
larvae to maturity on it. This plant was not available to the
author. It may have been possible to secure additional Pteryxia
plants which also grow on the slopes at the South Rim of the
Grand Canyon but, in view of the distance and time involved.
177
178
RONALD S. WIELGUS
/. Res. Lepid.
it was determined expedient to induce the larvae to accept still
another member of the Umbelliferae as a substitute host.
An attempt to reconstitute half of the remaining Pteryxia
plants by soaking in water for several hours was not successful.
The remains of the other half of the original host were then
placed in a small cooking pan holding approximately one pint
of water, which was then heated and brought to a boil and then
allowed to simmer for five minutes. The resulting solution was
then poured into glass jelly jars, capped and allowed to cool.
After cooling, this solution was used to water individual potted
plants of young (less than one foot high) Fennel (Foeniculum
vulgare). The watering was maintained on an hourly basis for
several hours, during which time the larvae were allowed to
find what little nourishment and moisture remained in the first
remaining half of the original host. The third-instar. No. 2, and
the two fourth-instar larvae. Nos. 3 and 4, moulted unobserved
prior to inspection on the morning of the seventh.
In the early morning of 7 June 1970 each larva was trans-
ferred to its individual potted Fennel plant in the hope of
obtaining acceptance. Larva No. 1 immediately accepted the
substitute host and fed periodically until the afternoon of the
eighth. Toward the end of that day it ceased feeding and re-
mained head downward on a petiole. It was determined that
this larva was preparing to moult.
The other larvae did not readily accept the Fennel and crawled
restlessly over the soil in the pots. Cut sprigs of fresh, tender
Fennel were then placed on the soil in each pot where the
larvae crawled, as it observed that the larvae experienced great
difficulty in attempting to crawl up the Fennel stems. Even with
this method the larvae nibbled but briefly on the Fennel tips,
which apparently did not completely satisfy their dietary re-
quirements, arxd continued their restless movements. The
feathery growth of the Fennel appeared to hamper the crawling
progress of the larvae and they continually lost footholds and
rolled over on their sides and backs.
On the morning of 8 June 1970 larvae Nos. 2, 3, 4 and 5 were
removed from the pots and placed in individual empty one
pound coffee cans. On the inside bottom of each can a cut-to-
fit disk of household paped towelling had been placed and on
this fresh sprigs of Fennel were laid. These sprigs were at first
obtained from the plants which had been watered with the
solution. The cans were then capped with the standard plastic
8(4):177-181, 1969(1970)
PAPILIO INDRA
179
lid that comes with each can and placed on a window sill away
from direct sunlight. This technique resulted in high humidities
inside the rearing cans but also served to prolong the freshness
of the foodplant. As the Fennel wilted in the course of time it
became necessary to replenish the rearing cans with fresh ma-
terial. This was done periodically during the day and on into
the evening hours. Each can was also emptied of accumulated
frass and a clean paper towel disk inserted. About 7:00 P.M.
(M.S.T. ) of the same day larva No. 4 began to feed earnestly
on the Fennel and continued for approximately twenty minutes.
This was in marked contrast with the earlier behavior which
exhibited rejection of the substitute host after several nibbles.
The two other fifth-instar larvae continued to maintain their
restlessness, pausing only occasionally to nibble, then resuming
their crawling. The fourth-instar larva. No. 2, remained quies-
cent and moulted unobserved early on the morning of the ninth.
On the morning of 9 June 1970 a review of the rearing cans
revealed that, of the three later fifth-instar larvae. No. 5 re-
mained motionless on its side on the bottom of the can in the
characteristic attitude assumed by Fapilio larvae prior to pu-
pation. This was confirmed by examination of the larva through
whose skin pupal features were distinguishable. It was noted
that this larva failed to spin the silken button and girdle so
characteristic of pupating larvae of this family. This may have
been due to the smooth metal side of the rearing can which
afforded little foothold for the larva. No difficulties were ex-
perienced, however, with similar rearing conditions for larvae
of Papilio cresphontes cresphontes Cramer, which simply spun
silken mats up a can’s side to their pupation sites. Of the other
two larvae. No. 4 continued feeding and No. 3 continued its
virtually ceaseless crawling. A stiff sheet of paper placed verti-
cally in the latter’s can did not elicit a response toward selection
of a pupation site and was ignored. The newly-moulted fifth-
instar larva. No. 2, accepted the Fennel and fed eagerly after
its mouthparts were sufficiently hardened. Larva No. 1 continued
to feed for a time after moulting but then contracted an unde-
termined ailment, evidenced by an expelling of a greenish liquid
from the mouth and excretion of a liquid frass. This larva rapidly
lost the ability to maintain a grip on the substitute host, dropping
to the soil and expiring shortly thereafter.
Larva No.^5 which had been determined to be prepupal was
placed in an upright tube of rolled stationery paper of slightly
180
RONALD S. WIELGUS
/. Res. Lepid.
longer length. On the morning of 10 June 1970 an examination
revealed that this larva had transformed to a chrysalis of slightly
smaller proportios (Emmel & Emmel, 1967), measuring 24 mm.
long by 8 mm. wide. This may have been brought about by a
reduced intake of nourishment in the last larval instar.
It was of interest to note that, with the exception of the ex-
pired No. 1 larva, the other larvae experienced great difficulty
in maintaining footholds and equilibrium on the Fennel sprigs.
This was not the case with larve of Papilio zelicaon Lucas which
the author has successfully reared on Fennel under similar rear-
ing conditions. Also of interest was the habit of kaihahensis
larvae of remaining quiescent for long periods of time, on the
order of a couple hours’ duration or more between feedings in
several instances, yet larval growth appeared to be rapid.
Feeding was noted to be avid in all stages, both on Pteryxia
and Fennel. Some difficulty was experienced by the larvae
feeding on the Fennel sprigs as the long, thin filaments continu-
ally slipped past their grip. A preference was shown for feeding
to begin at the terminal portion of each filament, though in some
instances the larvae would nip off the filaments mid-way and
feed upon the cut-off portions by holding these with their true
legs. After the larvae had fed several times upon the Fennel
sprigs from the plants watered with the solution, they were given
fresh sprigs from untreated plants. These were accepted without
hesitation. From then on only sprigs from untreated plants were
offered to the larvae.
At 10:09 P.M. (M.S.T. ) larva No. 3, which had previously
exhibited the most reluctance to feed, accepted the Fennel and
proceeded to feed avidly for approximately 10 minutes, where-
upon feeding terminated and did not resume again. Symptoms
of the ailment noted with larva No. 1 were exhibited by this
larva at 6:15 P.M. (M.S.T.) on 11 June 1970. The larva gradu-
ally lost mobility and slowly shrank in size during the next
several hours. However expiration, which appeared to be caused
by a combination of starvation and dehydration, did not occur
until 12 June 1970, probably due in part to the larger size of
this larva. At 7:30 A.M. (M.S.T.) of that day larva No. 4 also
excreted a voluminous liquid frass but did not exhibit the fatal
symptoms previously noted in the other larvae. Instead it pro-
ceeded to fashion a silken mat on the side of the rearing can
prior to assuming the pupation position and at 9:45 P.M.
(M.S.T.) it slipped into the silken girdle. Pupation took place
8(4):177-181, 1969(1970)
PAPILIO INDRA
181
unobserved during the early morning hours of the fourteenth.
This chrysalis measured 29 mm. long by 9 mm. wide.
Larva No, 2 continued to feed on the Fennel until 15 June
1970. Prior to selecting a pupation site, in this case on the screen
cover which replaced the plastic lid when the larva ceased feed-
ing, this larva, too, excreted a voluminous liquid frass and soiled
the paper towel disk extensively. It is not known at this time
whether defecation of a liquid frass at larval maturity is 'the
rule with this species or is caused by feeding on Fennel. This
larva pupated at 11:30 P.M, (M.S.T. ) on 16 June 1970 and a
perfect adult female eclosed prior to sunrise on 28 June 1970.
However, the chrysalis of larva No. 4, which had developed
to the verge of eclosion, died of unknown causes on 26 June
1970. At the time of this writing, 28 June 1970, the chrysalis
produced by larva No. 5, though still viable, shows no signs of
development and may have entered diapause.
It is interesting to speculate on the possibility of selective
breeding utilizing larger numbers of larva in order to develop
Fennel feeding populations. Since Fennel is easily grown from
seed, such host acceptance would offer wider study by serious
workers of the biology of this member of the indra complex.
The small sample combined with high mortality did not permit
the author to pursue this facet of his rearings.
ACKNOWLEDGMENT
The author would like to thank Dr. Frank F. Hasbrouck, As-
sociate Professor of Zoology and Curator of Insects, Arizona
State University, for critically reviewing the manuscript.
LITERATURE CITED
EMMEL, THOMAS C., and JOHN F. EMMEL, 1967. The biology of
Papilio indra kaibabensis in the Grand Canyon. Jour. Lepid. Soc.
21: 41-49.
POSTSCRIPT
The chrysalis produced by larva No. 5 on 10 June 1970,
which indeed had entered diapause, eclosed a perfect female
at dawn on 25 September 1970, Diapause was terminated by
refrigerating the chrysalis for thirty days, from August 4 to
September 4, then removing from refrigeration and maintain-
ing at room temperature until eclosion. During the period prior
to eclosion humidity was provided by placing the paper cylinder
containing the chrysalis on a water-moistened paper towel.
Journal of Research on the Lepidoptera
8(4): 182, 1969(1970)
1160 VV Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
HABITAT “ Colias philodice eriphyle
and Colias eury theme
W. HOVANITZ
These two species of Colias co-inhabit certain locales rang-
ing from the Sierra Nevada of California to the Atlantic Ocean.
Where the habitats coincide, there is extensive hybridization
of the two mutually fertile species. The locality shown here
is the Round Valley of Inyo and Mono counties, California, at
an elevation of about 4,000 feet, late June, 1970. In the back-
ground are the White Mountains, directly north, with White
Mtn. peak at about 14,500 feet. This is the most southern
locality known for C. philodice in California though in the
past (1920s) it was known as far south as Olancha.
182
Journal of Research on the Lepidoptera
8(4):183^186, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
EUPHYES DUKESI AND OTHER
ILLINOIS HESPERIIDAE
RODERICK R. IRWIN'
24 East 99th Place, Chicago, Illinois
For the past several years, Dr. John C. Downey and I have
been engaged in a survey of the butterflies of Illinois, which is
expected to lead to the publication of an annotated checklist of
the butterflies of that state. During the course of this study,
numerous interesting records of Hesperiidae have been obtained,
some of which I believe justify publication separately and in
advance of the larger work, since they represent significant range
extensions of the species involved.
Euphyes dukesi (Lindsey). Of perhaps the greatest interest
and importance has been the fact that this intensely local
skipper is apparently established in southern Illinois. Previously
published records were summarized by Mather (1963; 1966).
Price and Shull ( 1969 ) have recently published the first record
of dukesi from Indiana. A specimen from Blackwater, Prince
Albert County, Virginia, VI-13-64, leg. John Bauer, in the Car-
negie Museum collection, represents an apparently previously
unpublished Virginia record.
A single male of this species is in the collection of the Illinois
Natural History Survey, Urbana, Illinois. It was taken at Karnak,
Pulaski County, in extreme southern Illinois, on September 2,
1924, by T. H. Prison. In addition to the data label, this speci-
men bears a label reading “Upper side like / Type - Atrytone /
dukesi / Lind. / underside / more contrasting / Det / B[aj'nes].
& B[enjamin].” This appears to have been the first record of
the species following its original description, and consequently
the earliest record from other than the type locality. Its exist-
ence has apparently remained unpublished until the present.
Downey took another male dukesi in the Pine Hills region of
Union County, in southern Illinois, on September 10, 1966. Mr.
H. A. Freeman kindly confirmed my identification of this speci-
men, which is much darker on the upper side than any other
dukesi I have seen. It is almost wholly black, with only the
faintest traces of fulvous basally on the fore wings and discally
on the secondaries.
^Honorary Curator of Lepidoptera, Illinois State Museum.
183
184
R. R. IRWIN
/. Res. Lepid.
I took a rather worn male and a perfect female dukesi be-
tween 4:15 and 4:45 P.M. on August 31, 1969, in the same area
where Downey found the species in 1966. The insects were
found in an opening near the edge of a deciduous forest, just
south of a road paralleling the south side of the Big Muddy
River, about a half mile east of Illinois highway 3. The ground
at this point was entirely dry, but there were extensive swamps
just east of the area. No additional individuals of the species
were seen; they may have been more plentiful within the swamp
proper, but the lateness of the hour prevented a search for them
there. This appears to have been one of the few reported in-
stances of the capture of dukesi on dry ground, and is of par-
ticular interest since Mather (1963) indicates that the butterfly
seldom strays from its marsh habitat.
Collecting by Downey and his associates in the Pine Hills and
Lusk Creek (Pope County) regions of southern Illinois, as well
as examination of museum collections, has revealed the presence
of other Hesperiidae which had not been expected to occur in
Illinois. Among these are three species of Amhlyscirtes which
to my knowledge have not previously been recorded from so far
north in the Mississippi Valley.
Amhlyscirtes Carolina Skinner. Two specimens from the Pine
Hills, Union County, September 1, 1966, leg. J. C. Downey. The
range of this species is given by Klots (1951) as “Georgia to
Virginia” and it was unreported from any other area until
Mather and Mather (1958) published a Mississippi record. This
record from Illinois represents a further range extension. Identi-
fication of this and the following species was confirmed by
Freeman.
A. aesculapius (Fabr. ). Specimens from Pine Hills, Union
County, August 21, August 25 and September 1, 1966, leg. J. C.
Downey, and from Lusk Creek near Eddyville, Pope County,
July 7, August 9 and August 30, 1967, leg. J. C. Downey.
A. linda H. A. Freeman. I found a specimen which appeared
to be linda in the series of A. belli H. A. Freeman in the collec-
tion of the Field Museum of Natural History. It was taken at
Makanda, Jackson County, on July 8, 1896, by A. J. Snyder. I
sent it to Freeman, who confirmed my identification. Like the
two preceding species, this is the northernmost record of linda
known to me.
A. belli H. A. Freeman. This species was itself taken by
Downey at the Lusk Creek locality near Eddyville, Pope County,
August 9, 1967, 1 ^1$. This species has previously been
8(4):183~186, 1969(1970)
EUPHYES DUKESI
185
recorded from St. Louis County, Missouri (Remington, 1956).
These specimens were also determined by Freeman.
Autochton cellus ( Bdv. & Lee. ) . This scarce and local species
was taken in the Pine Hills region on July 26, August 21 and
September 1, 1966 (collector not indicated).
Hesperia ottoe Edw. Nielsen (1958; 1960) described the dis-
covery of this species in Michigan, and stated that this repre-
sented its first reported occurrence east of the Great Plains. The
wide gap between these two areas is partially bridged by the
finding of ottoe in Illinois. A female of this species was taken
by Alex K. Wyatt at Waukegan, Lake County, on July 28, 1946.
Thomas Taylor has taken a number of specimens at Mason
State Forest, Mason County, on the following dates: July 19,
1963, 1 $ ; July 11, 1964, 1 $ ; and July 19, 1964, 8 ^ 3 ? . The
first-named specimen was examined by H. A. Freeman, who
confirmed its identity. The others agree with it exactly. It
may perhaps be significant that the record from Waukegan
was six years earlier than the first Michigan record given by
Nielsen.
The preceding species discussed have not previously been
recorded from Illinois. The two following have been; but addi-
tional comment on their occurrence in that state may be worth-
while here.
Problema hyssus (Edw.). The first record of hyssus from
Illinois was given by Remington (1956) from Elsah, Jersey
County. Since then it has been found at five other locations in
the state. They are Peoria, Peoria County; Mason State Forest,
Mason County; Streator, La Salle County; Valmeyer, Monroe
County, and Perryton Township, Mercer County. Complete
data of capture will be given in the “Butterflies of Illinois,” in
preparation. It is interesting and perhaps significant that all
these localities lie at no great distance from either the Illinois
or Mississippi Rivers.
Thymelicm Uneola (Ochs. ). Since the publication of my paper
on this adventive European species in Illinois ( Irwin, 1968 ) , only
a single additional record of the skipper from outside the Chi-
cago metropolitan area has been reported: Divine, Goose Lake
Township, Grundy County, VI-22-68, leg. P. J. Conway. This
locality is almost exactly halfway in a direct southwestwardly
line between the Chicago area and Streator, La Salle County.
186
R. R. IRWIN
J. Res. Lepid.
The latter, therefore, remains the point of greatest range exten-
sion in Illinois from the Chicago area where it was first ob-
served. This is somewhat surprising in view of its rapid spread
heretofore; but it may reflect lack of collecting and reporting
rather than absence of the species in a wider area.
Intensive collecting of skippers in Illinois, particularly in the
southern portion of the state, may well yield results of equal
interest and significance to those presented in this paper. It is
unfortunate that so many amateur collectors neglect this fasci-
nating group as difficult, uninteresting, or both!
Specimens on which the records in this paper are based are
in the collections of the Illinois Natural History Survey, Southern
Illinois University, the Field Museum of Natural History, and
the personal collection of Mr. Thomas Taylor, of Peoria, Illi-
nois.
I am grateful to Mr. H. A. Freeman for the determinations
which are ascribed to him herein.
LITERATURE CITED
IRWIN, R. R., (1968). Thymelicus lineola ( Hesperiidae) in Illinois. Jour.
Lepid. Soc. 22:21-26.
KLOTS, A. B., ( 1951). A field guide to the butterflies. 349 pp. 40 pi. 8 fig.
Houghton Mifflin Co., Boston.
MATHER, B., (1963). Euphyes dukesi, a review of knowledge of its dis-
tribution in time and space and its habitat. J. Res. Lepid. 2:161-169.
(1966). Euphyes dukesi — Additional record. J. Res. Lepid. 5:254.
and K. MATHER, (1958). The butterflies of Mississippi. Tulane
Stud. Zool. 6:63-109.
NIELSEN, M. C., (1958). Observations of Hesperia pawnee in Michigan.
Lepid. News 12:37-40.
(I960). A correction on Hesperia pawnee in Michigan. Jour.
Lepid. Soc. 14:57.
PRICE, H. F. and E. M. SHULL, (1969). Uncommon butterflies of north-
eastern Indiana. Jour. Lepid. Soc. 23:186-188.
REMINGTON, P. S., (1956). Some observations on the Hesperiidae of the
St. Louis area. Lepid. News 9:190-195.
Journal of Research on the Lepidoptera
8(4):187-193, 1969(1970)
1160 W. Orange Grove Ave., Arcadia, California, U.S.A. 91006
© Copyright 1969
HESPERIA METEA LIFE HISTORY STUDIES
(HESPERIIDAE)'
J. RICHARD HEITZMAN" and ROGER L. HEITZMAN
3112 Harris Ave., Independence, Missouri
Hesperia metea Scudder ranges widely over the eastern half
of the United States. There is clinal graduation from typical
H. metea found in the New England states to the much darker
and larger H. metea Ucinus (Edwards) of eastern Texas. Speci-
mens from the Ozark plateau region of Missouri and Arkansas
are slightly smaller than Ucinus but otherwise compare well with
that population. This is a univoltine species with imagines flying
in any given region for a few weeks in the spring. The typical
habitat in the Ozark region is found on dry, often rocky hillsides
in direct proximity to woodland areas. Beard grass {Andropo-
gon gerardi Vitm. ), a characteristic plant of the Ozark flora,
serves as the larval host. H. metea is one of the earliest native
spring species, adults emerging with the flowering of red bud
and wild plum trees. The wary males are found resting on
bare patches of earth or visiting early flowers. Bird's-foot violet
(Viola pedata) and wild strawberry (Fragaria virginiana var.
illinoensis) are especially attractive. Females are not as wild
and can be observed flying about the larval host plants where
they settle near the base of the plants and crawl among the
dried leaves and litter laying eggs. Since females fly a little
later in the season they express some additional flower pref-
erences and frequent wild larkspur (Delphinium carolinianum) ,
wild hyacinth (Camassia scilloides), and Verbena species.
This species is the possessor of an interesting and unusually
complicated life cycle. Females lay freely in captivity with or
without the presence of Andropogon. During the first few
instars the larvae are nocturnal in feeding habits: remain-
ing hidden in their tents during daylight hours. In the later
instars the larvae live deep within the base of the plants:
actually tunneling below ground level. During the hot weather
of late July, August, and early September the larvae spend long
^Contribution No. 174, Bureau of Entomology, Division of Plant Industry, Florida De-
partment of Agriculture and Consumer Services.
^Research Associate, Florida State Collection of Arthropods, Division of Plant In-
dustry, Florida Department of Agriculture and Consumer Services, Gainesville.
187
Fig. 1.— Hesperia metea Scudder, 1-2, Head of final instar larva, frontal
and left lateral aspect. 3, Ovum. 4, Head of first instar larva, frontal aspect.
5, Mature larva.
188
HEITZMAN AND HEITZMAN
J. Res. Lepid.
8(4);187-193, 1969(1970)
HESPERIA METEA
189
periods in aestivation hidden deep within their tunnels. The
larvae are fully developed when the first cold weather of fall
arrives which provides the stimulus for hibernation. The hiber-
nation chamber is constructed between two or more grass
blades deep in the center of the grass plant. The chamber is
thickly lined with silk and tightly sealed. Pupation supposedly
occurs with the first warm days and rains of early spring. We
have reared this pesky species from ova to hibernating larvae six
different years but have yet to obtain a single pupa. We have
tried numerous indoor and outdoor arrangements including
enclosing entire growing plants in the garden with screen wire
cages. H. metea does not occur in the Independence area and
we have had no opportunity to attempt rearing in its native
Ozark haunts. There may be edaphic problems involved since
climatic conditions are essentially the same in both areas.
Many hours have been spent in the field during early spring
looking for the “needle in the haystack.” By carefully pulling
apart the dried Andropogon clumps we have found pupae of
Atrytonopsb hianna (Scudder), Everes comyntas (Godart),
Apantesis anna Grote, and seven species of Noctuidae. At least
a dozen m-etea hibernation chambers with the shriveled remains
of their occupants have been found. This suggests that the
natural mortality rate may be high during this dormant period.
The following descriptions, minus the elusive pupa, are based
upon six rearings from ova to hibernating larvae and many field
observations conducted in the vicinity of Warsaw, Missouri,
and Fayetteville, Arkansas. The illustrations were drawn by the
junior author from specimens collected near Warsaw, Missouri
during 1968 and 1969.
OVUM: Width 1.50mm, Height 1.25 mm. Creamy white, no
visible markings. Eclosion in seven to eight days. Micropyle
darkens on fifth day in fertile ova.
FIRST INSTAR LARVA: Head deep glossy purple, thinly cov-
ered with short pale setae. Pr ©thoracic shield deep purplish
black. Body white, unmarked, sparsely covered with white setae,
some longer hairs on anal segment. The emerging larvae eat
from one half to an entire egg shell. After eating the egg shell
the larvae make a narrow open tent along a leaf edge a few
inches from the tip. Small notches are eaten from one side of
the grass blade for several inches up and down the leaf includ-
ing the tent itself. After a few days of feeding the tent is en-
larged and a greater amount of silk expended than for the
initial structure. On the second day of feeding the body as-
sumes a slight greenish tint. Stadium period: seven to nine days.
190
HEITZMAN AND HEITZMAN
J. Res. Lepid.
2. — Hesperia metea Scudder, 1, Adults male and female, dorsal and
ventral view. 2, Setae of first instar larva, prothorax, mesothorax and eighth
abdominal segment, all in left lateral aspect. 3, Setae of suranal plate,
dorsal aspect.
8(4):187-193, 1969(1970)
HESPERIA METEA
191
SECOND INSTAR LARVA; Head deep purplish black, granu-
lose, thickly covered with short white setae. Prothoracic shield
purplish black. Body pale greenish white, the three posterior
segments paler. Body thinly covered with short white setae,
some longer hairs curving back from anal segment. A few
partial tents are constructed during this instar but in most cases
the larvae hide at the base of the leaves in a fold of the leaf
when not feeding. On the last day they spin a silk covering and
molt within this protection. Stadium period: 19 to 21 days.
THIRD INSTAR LARVA: Head, prothoracic shield, and first
pair of thoracic legs reddish purple. Head thickly covered with
short pale setae, mandibles black. Two pale orange areas visible
low on front of head capsule between stemmata and laterofacial
suture lines. Face deeply cleft at midcranial inflection which is
black with narrow orange edging. Body pale creamy gray, in-
tersegmental folds pale yellow. Body thickly covered with
minute black setae. Anal spiracles marked by a black dot.
Stadium period: 10 to 12 days.
FOURTH INSTAR LARVA: Head deep reddish purple, deeply
cleft at midcranial inflection which is edged with deep orange.
A large orange area is present between the stemmata and
laterofacial suture lines. Thoracic legs black tipped. Body
creamy gray with tiny pale orange setae, intersegmental folds
pale orange yellow. First thoracic and two anal spiracles marked
by black dots. Prothoracic shield black. Stadium period: 10
to 12 days.
FIFTH iNSTAR LARVA: Head deep reddish purple, granu-
lose, covered with short orange setae, mandibles and stemmata
black. Midcranial inflection edged by narrow orange lines, Irons
pale cream color. A small orange spot is located on each side
between the stemmata and the laterofacial suture lines. Pro-
thorax white, conspicuous. Prothoracic shield shiny black.
First thoracic and anal spiracles marked with a large black dot,
a tiny black dot at other spiracles. Body grayish orange, ab-
dominal area paler, thickly covered dorsally with minute orange
setae, a few longer hairs on anal segment, intersegmental folds
paler. First two pair of thoracic legs deep purple, last pair
pale brown. Integument opaque with a leathery texture. Larvae
in this instar feed voraciously for about a week after which they
become restless and leave the host plant. After wandering about
for a day they begin spinning thinly lined silken tubes one to
two inches in length in the center of the host plants near ground
level. Very little is eaten for the next three weeks, only a few
notches here and there over the plants. Every few days the
larvae move to new spots and start a new tube. The larvae are
192
HEITZMAN AND HEITZMAN
/. Res. Lepid.
extremely nervous during this period. Even approaching the
plants causes them to move uneasily and may have been the
cause of frequent moves to new quarters. This aestivation
period is apparently brought on by the dry midsummer weather.
The size of the larvae during the last two weeks of this instar
remains nearly constant. Larvae being reared outside were
spurred to prepare for and enter the sixth instar after summer
showers had fallen. Larvae being reared indoors were stimu-
lated by repeated soakings of rainwater. Since the rainfall was
the apparent factor governing the stadium period of this and
the next (sixth) instar the duration time varied greatly: from
19 to 31 days in the fifth instar to a maximum of 51 days in
one instance in the sixth instar. After the moisture stimulus a
period of several days of heavy eating would begin followed by
rapid molting and ingress into the next stadium period.
SIXTH INSTAR LARVA: Head deep brownish purple, granu-
lose, mandibles deeper purple. Midcranial inflection bordered
with bright orange lines. There is a duller orange area between
the stemmata and laterofacial suture lines with a small extension
rising vertically opposite the midcranial inflection. A small
orange raised area is located directly posterior to the stemmata.
Prothorax shiny white, prothoracic shield jet black. Body an
unusual pinkish gray best described as grayish flesh, abdomen
and prolegs pale flesh color, anal segment paler dorsally, almost
translucent. First pair of thoracic legs black, posterior pair pale
brown. Integument semi-transparent, dark areas inside body
showing as blurred undulating spots. Heart line visible as a
dark, pulsing middorsal line. Intersegmental folds dark pink,
smooth in appearance. Small white setae visible over the body,
more noticeable on anal segment. Aestivation occurs off and on
during the sixth instar with the larvae retiring to their silken
lined tubes deep within the base of the host plants. Sometimes
several days elapse without any noticeable evidence that they
have emerged. At other times the larvae become restless and
wander about over the plants eating small notches here and
there. The larvae require three days preparation before molting.
Stadium period varies greatly and is seemingly dependent upon
the arrival and amount of moisture received.
SEVENTH (FINAL) INSTAR LARVA: Length of mature
larvae is 31 to 34 mm. Body grayish brown with slight lavander
overcast, abdomen and prolegs slightly paler. Integument slight-
ly translucent with a wrinkled appearance between interseg-
mental folds. Prothorax white, prothoracic shield and thoracic
legs jet black. Spiracles marked by black dots. Head dark
purple with orange lines paralleling midcranial inflection.
8(4):187-193, 1969(1970)
HESPERIA METEA
193
Orange lines parallel laterofaciai suture lines and enter a paler
cream colored area between the stemmata and laterofaciai
sutures, this pale area with an uneven vertical extension. A
protruding orange area is located just posterior to the stemmata
of which three is largest, two and ' four about equal in size, one
and six equal and five the smallest. Stemmata positioned as in
sketch of head capsule. Extent and intensity of head markings
is variable with different specimens tending to become obscure
near end of final instar. The illustrations of the head capsule
markings are from specimens that have just entered the final
instar when they are sharp and clear. The larvae are lethargic
during the final instar. When disturbed they will curl into a
tight ball and feign death, remaining thus for long periods of
time, as long as 35 minutes by actual count. They feed leisurely
during the day in the open, retiring to their silk lined tubes
when not feedng. The final tube tent is constructed in the
center of the plants, extending two or three inches into the base
of the plant. The final instar larvae have two fluffy white areas
of waxlike flakes beneath the posterior segments of the abdomen.
In other cases where we have observed these wax flake patches
on larvae the pupae were subsequently found coated with them
(perhaps an excess moisture repellent since in at least one
case, Euphyes dion Edwards, the pupae are occasionally sub-
merged under water for lengthy periods ) . Stadium period quite
variable, hibernation being stimulated by cool weather which
occurs in late September in the Ozark region during normal
years.
We wish to express our thanks to Dr. Leo J. Paulissen, Fayette-
ville, Arkansas for valuable field assistance. We are indebted
to Dr. Alexander B. Klots, American Museum of Natural His-
tory and Dr. Howard V. Weems Jr., Florida State Dept, of
Agriculture for reading the manuscript and offering valuable
advice. We also owe our thanks to Dr. John R. Reeder, Yale
University for plant determinations concerning this and other
life history studies in progress.
Journal of Research on the Lepidoptera
8(4):194, 1969(1970)
1160 W. Orange Grove Ave.» Arcadia, California, U.S.A. 91006
© Copyright 1969
HABITAT — Oeneis chryxus Stanislaus
W. HOVANITZ
Shown here is the type locality for the race Oeneis chryxus
Stanislaus on the ridge at Sonora Pass, California, elevation
9,700 feet, late June, 1970. Both north and south from this
point, the ground color of the wings becomes lighter, termi-
nating in the disjunct race Oeneis chryxus ivallda along the
crest of the Sierra Nevada. To the east in the Sweetwater
Range, the color remains brown. Oeneis chryxus ivallda has
the distinction of being separated into two parts, isolated
from one another by this intrusion of “brown” genes.
194
BOOKS RECEIVED
NORDSTROM, FRITHIOF, SVEND KAABER, MAGNE OPHEI M, and
OLAVI SOTAVALTA
DE FENNOSKANDISKA OCH DANSKA NATTFLYNAS UTBREDNING
(Distribution of the Macro Lepidoptera of Fennoscandia and
Denmark - Noctuidae) . Lund, 1969,
CONEL, J, L.
LIFE AS REVEALED BY THE MICROSCOPE
Philosophical Library, New York, 1969 ^7. 95
UNESCO
STUDY ABROAD XVIII
United Nations Educational, Scientific and Cultural Organization
Paris, 1969 ^6. 00
CURTIS, WILLIAM
A SHORT HISTORY OF THE BROWN- TAIL MOTH
Curwen Facsimile, Plaistow, 1969
Entomological Reprint Specialists ^9. 30
P. O. Box 207
East Lansing, Michigan 98823
SCHMID, MICHAEL and BRADFORD M. ENDICOTT
MARIPOSAS DE VENEZUELA
Nordgrafik Ltd. , Copenhagen, Denmark, 1968
Entomological Reprint Specialists ^9, 95
P. O, Box 207
East Lansing, Michigan 48823
COMMON, I. f: B,
AUSTRAILIAN BUTTERFLIES
Jacaranda Press Pty. , Brisbane, Australia, 1964
BLANDINO S. J. , GIOVANNI
THEORIES ON THE NATURE OF LIFE
Philosophical Library, New York, 1969 ^6. 00
SAVORY, T. H.
INTRODUCTION TO ZOOLOGY
Philosophical Library, New York, 1968 ^6. 00
MOUCHA, JOSEF
BUTTERFLIES
The Hamlyn Publishing Group LTD. , Middlesex, 1968
FORBES, WILLIAM T. M.
THE LEPIDOPTERA OF NEW YORK AND NEIGHBORING STATES
Part 1
Entomological Reprint Specialists, East Lansing, Michigan, 1969
^17. 50
BOOKS RECEIVED [continued]
DARLINGTON JR., PHILIP J.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD
McGraw-Hill Book Co. , 1968 ^2. 95
MOSHER, Ph. D. , EDNA
LEPIDOPTERA PUPAE
Entomological Reprint Specialists, East Lansing, Michigan,
1969 ^9. 95
CLARK, GOWAN C. and C. G, C. DICKSON
SOME SOUTH AFRICAN BUTTERFLIES
Longmans Green and Co. , Cape Town, 1952
LESTER, JAMES D.
WRITTING RESEARCH PAPERS
Scott, Foresman and Co., Glenview, 111., 1967
LEYTE-SAMAR STUDIES Vol. II, No. 1, 2; Vol. IV, No. 1
DWU Graduate School Publication, Tacloban City, Philippines
1968, 1970
NOTICES
BOOKS:
BUTTERFLIES. A concise guide in colour, Josef Moucha, ill. by
Vlastiinil Choc. Paul Hamlyn, Hamlyn House, The Centre,
Feltham, Middlesex. G.B.
BIOGEOGRAPHY OF THE SOUTHERN END OF THE WORLD.
Philip J. Darlington, Jr. McGraw Hill paper hack reprint, N.Y.
THEORIES ON THE NATURE OF LIFE. Giovanni Blandino, S.J.
Philosophical Library, N.Y.
INTRODUCTION TO ZOOLOGY. Theodore H. Savory. Philosophical
Library, N.Y.
WANTED:
Brephklium exilis, B. fea, B. isophthalma. Life material and specimens
for distribution study. Roy Jameson, 2429 Wordsworth, Houston,
Texas 77025.
ARGYNNIS. Local and world wide, for world biogeographic study.
Also related forms under whatever name. William Hovanitz, 1160
W. Orange Grove Avc., Arcadia, California 91006.
IN PREPARATION:
BUTTERFLIES OF NORTH AMERICA. William Hovanitz. Illustrat-
ing in color all the species and races of butterflies of the Nearctic
region. Will be the first book on butterflies to use the New
Systeniatics, biogeographical and genetic approach to an under-
standing of this group of insects.
NEEDED:
Manuscripts for immediate publication in this JOURNAL. With color
n^ay be delayed; black and white immediate. Needed to bring our
schedule up-to-date.
TO SAVE WORK FOR THE EDITOR please write notices on a
3 X 5 card in the form desired and they will be printed in the
next following issue of the JOURNAL.
Volume 8
Number 4
December, 1969
IN THIS ISSUE
f \
^ \ \
Seasonal changes in organization of
tropical rain forest butterfly populations
in Panama.
Thomas C. Emmel and Charles F. Leek 133
Studies on Nearctic Euchloe. Part 6.
Systematics of adults. Paul A. Opler 153
Laboratory production of the Monarch
Butterfly, Danaus plexippus.
F. Munger and T. T. Harriss 169
The rearing of Papilio indra kaibabensis,
Ronald S. Wielgus 177
Habitat — C alias philodice eriphyle
and Colias eurytheme. William Hovanitz 182
Euphyes dukesi and other Illinois Hesperiidae.
Roderick R. Irwin 183
Hesperia metea life history studies.
J. R. Heitzman and Roger L. Heitzman 187
HabiioX—Oeneis chryxus Stanislaus.
W. Hovanitz 194
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